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1/21999 Macmillan Magazines Ltd
Women prefer slightly feminized malefacial shapes1. Such faces
(Fig. 1a) are
given positive personality attributions1 thatmight correlate
with actual behaviour2. Incontrast, masculine features seem to
sig-
nal immunological competence
3
. Heritablebenefits can be realized only if conceptionfollows
copulation, so women might bemore attentive to phenotypic markers
indi-cating immunological competence duringthe follicular phase of
the menstrual cyclewhen conception is most likely4,5. Consis-tent
with this hypothesis is the observationthat womens preference for
the odour ofmen with low fluctuating asymmetry (acorrelate of
testosterone-facilitated trait sizeand developmental stability)
increases withthe probability of conception across themenstrual
cycle5. Symmetrical men reportmore extra-pair copulation partners6,
and
extra-pair copulation rates peak in mid-cycle7. Here we show
that female preferencefor secondary sexual traits in male
faceshapes varies with the probability of con-ception across the
menstrual cycle.
Japanese subjects reporting regular men-strual cycles and no use
of oral contracep-tives (n39, mean age 21 years, mean cyclelength
30 days) answered questions aboutthe average cycle length and date
of onset ofprevious menses. From this information,two testing
sessions were arranged for thesampling of individuals preferences
duringphases of both high and low conception risk
in their menstrual cycles. Ovulation wasassumed to occur 14 days
before the onsetof menses. Sessions after ovulation andbefore the
onset of menses (the lutealphase) and sessions during menses
wereclassified as low conception risk. Sessionsin the follicular
phase (between the end ofmenses and ovulation) were classified
ashigh conception risk4,5,7. Subjects wereasked to select the face
that they consideredmost physically attractive (miryoku-teki)from
five Caucasian and, separately, fiveJapanese male faces (40% and
20% femi-nized, on average, and 20% and 40% mas-culinized).
Subjects were also asked whether
they currently had a steady boyfriend(tsukiatteiru). Stimuli
were symmetricalversions of those used (and illustrated) inprevious
research1 but with hair visible, pre-sented with full
counterbalancing.
Repeated-measures analysis of variance(ANOVA) showed a
significant main effectof conception risk (variance
ratio(F(1,37))9.47; P
8/3/2019 Menstrual cycle alters face preference
2/21999 Macmillan Magazines Ltd
partner whose low masculine appearancesuggests cooperation in
parental care(long-term preferences are unchangedacross the
menstrual cycle) but occasionallycopulate with a male with a more
masculineappearance (indicating good immunocom-petence) when
conception is most likely.Sexual behaviour arising from cyclic
prefer-ences might allow individuals to accrue
benefits from polyandry while maintainingthe advantage of
ostensive monandry.I. S. Penton-Voak*, D. I. Perrett*,
D. L. Castles, T. Kobayashi, D. M. Burt*,L. K. Murray*, R.
Minamisawa
*School of Psychology, University of St Andrews,
Fife KY16 9JU, UK
e-mail: [email protected]
Hasegawa Laboratory,
Department of Life Sciences, University of Tokyo,
Komaba, Tokyo 153, Japan
School of Life Sciences,
Roehampton Institute London, Whitelands College,
London SW15 3SN, UK
1. Perrett, D. I. et al. Nature394, 884887 (1998).
2. Berry, D. S. & Wero, J. L. F.J. Pers. 61, 497523
(1993).
3. Folstad, I. & Karter, A. Am. Nat. 139, 603622 (1992).
4. Regan, P. C. Can. J. Hum. Sexual. 5, 145156 (1996).
5. Gangestad, S. W. & Thornhill, R. Proc. R. Soc. Lond. B
265,
927933 (1998).
6. Gangestad, S. W. & Thornhill, R. Evol. Hum. Biol. 18,
6988
(1997).
7. Baker, R. R. & Bellis, M.A. Human Sperm Competition:
Copulation, Masturbation and Infidelity(Chapman & Hall,
London, 1995).
8. Christenfeld, N. & Hill, E. A. Nature378, 669 (1995).
9. Frost, P. Percept. Motor Skills79, 507514 (1994).
10.Grammer, K. Ethol. Sociobiol. 14, 201208 (1993).
11.Graves, J., Ortegaruano, J. & Slater, P. J. B. Proc. R.
Soc. Lond. B
253, 37 (1993).
both indicated that the juxtaposition of cellsthat did and did
not express Rfng is suffi-cient for the formation of the apical
ecto-dermal ridge (AER), an organizing centrethat forms at the
boundary between dorsaland ventral cells at the distal edge of
thelimb bud4,5. We have used homologousrecombination in mouse
embryonic stem
cells to investigate Rfng expression in themouse limb bud and to
test whether it isnecessary for the formation of the AER.
We inserted the Escherichia coli lacZgene into the Rfnggenomic
locus to createthe RfnglacZKI knock-in allele. Analysis
of-galactosidase activity in RfnglacZKI limb budsrevealed
expression in the developing andmature AER (Fig. 1ac). To create a
nullallele, Rfng1-8neo, we deleted 2.4 kilobases ofthe Rfng genomic
locus containing theentire coding region (exons 18) andreplaced it
with a PGKneo-selection cas-sette (which confers a resistance to
the
antibiotic neomycin and acts as a means forselecting these
clones) in the opposite tran-scriptional orientation.
No homozygous Rfng1-8neo-deficientmouse survived past birth
(n244 progenyfrom heterozygote intercrosses on a mixed129P2C57BL/6J
hybrid background;n70 progeny from heterozygote inter-
crosses on a mixed 129P2129S6 hybridbackground). The mutant mice
displayedseveral developmental defects, including acleft palate and
reduced mandible (64%),exencephaly (24%), and forelimb defects(12%)
consisting of soft-tissue syndactylyor oligodactyly (total n225 on
a mixed129P2/OlaHsdC57BL/6J hybrid back-ground, examined between 18
days post-coitum (d.p.c.) and birth; Fig. 1).
We went on to remove the PGKneo-selection cassette to create the
Rfng1-8 nullallele by two independent methods: matingRfng1-8neo
mice to mice that expressed Cre
scientific correspondence
742 NATURE | VOL 399 | 24 JUNE 1999 | www.nature.com
Limbs move beyond theRadical fringe
The fringe genes and the Notch signallingpathway are important
in limb develop-ment in vertebrates and in Drosophila, help-ing to
establish the organizing centres thatare required for the
proximaldistal out-growth of appendages15. Three vertebrateFringe
genes have been identified: Manic(Mfng), Lunatic (Lfng) and Radical
fringe(Rfng)48. Here we show that the mouseRfnggene is not required
for limb develop-
ment, even though it is expressed in thedeveloping limb bud. But
we have foundthat several developmental defects, which atfirst we
attributed to a loss ofRfngfunctionafter mutating the gene, in fact
arose as aresult of the insertion of a selection cassetteinto the
Rfnglocus that affects the expres-sion of a neighbouring gene or
genes. Ourfindings highlight the need for optimizationin designing
mutant alleles to probe devel-opmental processes9,10.
Studies ofRfngexpression in chick limbmutants and
gain-of-function experimentsin limb buds using retroviral
misexpression
p m
a b c
d e f
g h i
j k l
RfnglacZKI
Rfng1-8neo/ Rfng1-8/
9.5 10.5 12.5
Rfng1-8neo+/+
D V
p pm m
Figure 1 Rfng is expressed in the mouse limb bud and is not
required for limb development. Rfngexpression
in RfnglacZKI heterozygote limbs, as shown by X-gal staining,
at: a, 9.5 d.p.c.; b, 10.5 d.p.c.; and c, 12.5 d.p.c.
Dorsal (D), left; ventral (V), right. Replacement of the Rfng
locus with a PGKneo-selection cassette results in
pleiotropic developmental defects. d, g, j, Wild-type fetuses;
e, h, k, Rfng1-8neo homozygotes; and f, i, l, Rfng1-8
homozygotes at 18.5 d.p.c. df, Skeletal staining of the
secondary palate. Anterior is to the right. d, The
palatal (p; indicated by arrows) and maxillary (m) shelves are
nearly fused. e, Rfng1-8neo homozygote palatal
shelves (p; white arrows) have not elevated towards the midline,
and the maxillary shelves (m) are wider
than in the wild type. f, Rfng1-8homozygote, showing normal
palatal (p) and maxillary (m) shelf positioning.
g, Wild-type fetus with closed cranium. h, Rfng1-8neo
homozygote, showing exencephaly. i, Rfng1-8 homo-
zygotes are indistinguishable from wild type (g). jl, Skeletal
staining of the forelimbs. Anterior is up, posterior
is down. j, Rfng1-8neo wild-type limb has five digits. k,
Rfng1-8neo homozygote, showing soft-tissue syn-
dactylism of digits 4 and 5 (arrow). l, Rfng1-8homozygotes are
indistinguishable from wild type (j).
