Lower olfactory threshold during the ovulatory phase of the menstrual cycle

Lower olfactory threshold during the ovulatoryphase of the menstrual cycle

Evelia Navarrete-Palacios a,b, Robyn Hudson c, Gloria Reyes-Guerrero b, Rosalinda Guevara-Guzman b,*

a Hospital Central Militar, Ciudad de Mexico, Instituto de Investigaciones Biomedicas, Universidad Nacional

Autonoma de Mexico, 04510 Mexico, D.F., Mexicob Departamento de Fisiologıa, Facultad de Medicina, Instituto de Investigaciones Biomedicas, Universidad

Nacional Autonoma de Mexico, Apartado Postal 70250, 04510 Mexico, D.F., Mexicoc Departamento de Biologıa Celular y Fisiologıa, Instituto de Investigaciones Biomedicas, Universidad

Nacional Autonoma de Mexico, 04510 Mexico, D.F., Mexico

Received 15 November 2002; accepted 8 April 2003

Abstract

We investigated whether olfactory detection threshold is correlated with phase of the

menstrual cycle. Three hundred and thirty-two women 13�/49 years old were tested once

during either the follicular, ovulatory, luteal or menstrual phase, and 15 women 20�/43 years

old were tested at each of these phases across one complete cycle. In three non-cycling control

groups subjects were each tested once; 83 post-menopausal women 47�/86 years old, 60 pre-

pubertal girls 5�/12 years old, and 183 men 17�/30 years old. Odor detection thresholds were

determined using sniff bottles containing �/log 9.5 to �/log 6.0 concentrations of amyl acetate

presented in ascending order. Thresholds differed significantly across the cycle and were

lowest during the ovulatory and highest during the menstrual phase. Thresholds for all control

groups were higher than for the cycling women during the ovulatory phase. The results

confirm that olfactory threshold is related to phase of the menstrual cycle and thus possibly to

hormonal state.

# 2003 Elsevier B.V. All rights reserved.

Keywords: Olfactory sensitivity; Hormonal state; Men; Girls; Post-menopause

* Corresponding author. Tel.: �/525-550-3587; fax: �/525-623-2241.

E-mail address: [email protected] (R. Guevara-Guzman).

Biological Psychology 63 (2003) 269�/279

www.elsevier.com/locate/biopsycho

0301-0511/03/$ - see front matter # 2003 Elsevier B.V. All rights reserved.

doi:10.1016/S0301-0511(03)00076-0

mailto:[email protected]

1. Introduction

Olfactory researchers have long been interested in a possible relationship between

olfactory function and the menstrual cycle, and particularly as indicated by changes

in odor detection threshold. At present, the weight of evidence suggests increased

sensitivity, at least for some substances, around the time of ovulation and/or the

mid-luteal phase (Le Magnen, 1952; Vierling and Rock, 1967; Mair et al., 1978; Dotyet al., 1981, 1982), including as measured by olfactory reaction time (Narita et al.,

1992) and event-related potentials (Pause et al., 1994, 1996). Some investigators,

however, have reported increased sensitivity during the follicular phase (Henkin,

1974), or around menses (Koster, 1968; Doty, 1976), while still others have reported

decreased sensitivity at menstruation (Le Magnen, 1952; Schneider and Wolf, 1955;

Good et al., 1976; Mair et al., 1978; Moriyama and Kurahashi, 2000), or have failed

to find reliable, cycle-dependent changes in olfactory sensitivity (Amoore et al., 1975;

Herberhold et al., 1982; Filsinger and Monte, 1986; Hummel et al., 1991; Kanamuraand Takashima, 1991), including in olfactory event-related potentials (Nghiemphu et

al., 1995).

Such discrepancies may be at least partly explained by differences in methodology

among studies, including the small sample sizes sometimes used and differences in

the time points of the cycle taken for comparison. Interpretation of findings is also

made difficult in some cases by the lack of parallel testing of non-cycling control

groups. In the present study it was therefore our aim to investigate the relationship

between the menstrual cycle and olfactory threshold by testing a large sample ofcycling women not taking oral contraceptives, together with three non-cycling

control groups*/post-menopausal women, pre-pubertal girls and young men.

2. Methods

The study proposal and test procedures were approved by the ethics committees of

the Faculty of Medicine, National University of Mexico, and the Central Military

Hospital, Mexico City. Subjects, or in the case of the girls, their parents, were

informed as to the purpose of the study before obtaining their voluntary consent to

participate.

2.1. Definition of the menstrual cycle

This was defined as the time between the first day of one menstruation and the

first day of the next, with the first day of menstrual bleeding taken as day 1. Themenstrual phase was defined as cycle days 1�/5, the follicular or estrogenic phase as

days 6�/14, and the luteal or progestagenic phase as days 15�/28 or beyond. The

menstrual phase was defined in a uniform manner because of uncertainty as to the

exact length of menstruation for each of the large number of women recruited at

different stages of the cycle in Group 1 (see below). As described for Groups 1 and 2

E. Navarrete-Palacios et al. / Biological Psychology 63 (2003) 269�/279270

below, the ovulatory phase was defined as the day midcycle when basal temperature

rose by at least 0.3 8C.

2.2. Subjects

Participants were recruited from the campus of the National University and from

the Military Hospital. All were healthy, non-smoking residents of Mexico City, none

was taking medication and none had used oral contraceptives for at least 6 months

prior to the start of the study. Subjects were instructed not to use perfumed toiletries

on the day of the experiment and not to drink alcohol or coffee or to eat garlic,

onions or other spicy foods the day before. The following groups were formed (Table1).

2.2.1. Group 1

Three hundred and thirty-two regularly cycling women 13�/49 years old whose

cycles ranged from 28 to 30 days (mean 28.8, SD 0.3) tested once during one of the

four principle phases of the menstrual cycle*/menstrual, follicular, ovulatory or

luteal. To obtain subjects for testing during the ovulatory phase, women recruited

close to midcycle were instructed to take their oral temperature at the same time each

morning, to enter the values on a score sheet, and to report for olfactory testing

when their temperature rose by at least 0.3 8C. Only subjects adhering strictly to the

measuring regimen and showing a clear rise in temperature were included in this,consequently small, category.

2.2.2. Group 2

Fifteen regularly cycling women 20�/43 years old whose cycles ranged from 28 to30 days (mean 29, SD 0.3) tested at each of the four principle phases of one

menstrual cycle*/menstrual, follicular, ovulatory, and luteal. For five women testing

started in the menstrual phase, for four women in the follicular, for two in the

ovulatory, and for four in the luteal phase. As for Group 1, subjects were instructed

Table 1

Composition of the experimental groups according to age

Groups N Mean age in years (SD) Age range in years

Cycling women tested once 332 22.1 (4.7) 13�/49

Menstrual phase 87 23.6 (4.8) 13�/49

Follicular phase 80 22.6 (4.7) 13�/45

Ovulatory phase 21 22.7 (4.7) 13�/43

Luteal phase 144 22.3(4.7) 14�/49

Cycling women tested repeatedly 15 26.2 (5.5) 20�/43

Post-menopausal women 83 67.4 (8.2) 47�/86

Pre-pubertal girls 60 8.3 (2.8) 5�/12

Young men 183 21 (4.5) 17�/30

Navarrete-Palacios et al., 2003.

E. Navarrete-Palacios et al. / Biological Psychology 63 (2003) 269�/279 271

to take their oral temperature at the same time each morning, to enter the values on a

score sheet, and to report for olfactory testing when their temperature rose by at

least 0.3 8C.

2.2.3. Group 3

Eighty-three post-menopausal women 47�/86 years old, at least 6 months after the

last menstruation, tested once.

2.2.4. Group 4

Sixty pre-pubertal, non-menstruating girls 5�/12 years old, tested once.

2.2.5. Group 5

One hundred and eighty-three men 17�/30 years old, tested once.

2.3. Test procedure

Testing was carried out in a quiet room at the Military Hospital between 10:00 and

13:00 h. Subjects were seated in a comfortable chair at a room temperature of 18�/

22 8C.

In a first step, designed to obtain a rough estimate of each subject’s sensitivity so

as to reduce the number of concentrations presented in the subsequent finer

determination of threshold, subjects were presented with an ascending series of eightconcentrations of amyl acetate (J.T. Baker, Mexico) in 100 ml polyethylene squeeze

bottles equipped with a fine plastic nozzle and containing a 30 ml solution of the

odorant in distilled water. Based on pilot testing, the concentrations we used ranged

from �/log 9.5 to �/log 6.0 presented in ascending half log concentrations. Subjects

were asked to hold each bottle under their nose, to squeeze it gently, and to respond

with yes or no according to whether they detected an odor or not. They were only

allowed to sample the content of each bottle once and with an interval of at least 45 s

between bottles. Amyl acetate was chosen because of its frequent use in olfactorystudies, because it is generally agreeable, readily available, soluble in water, and

because its smell resembles banana and so might be considered familiar and easy to

detect for the population tested.

In a second step, a finer estimate of detection threshold was obtained by

presenting an ascending series of concentrations using a four-alternative forced-

choice procedure (cf. Koelega and Koster, 1974; Koelega, 1994). Subjects were

presented with a set of four bottles, two of which contained amyl acetate one dilution

step below the concentration the subject had detected previously, and two whichcontained 30 ml distilled water. Subjects were asked to identify the two bottles that

contained the odorant, the probability of achieving this by chance being only 17%

compared to 33% in the commonly used three-bottle forced-choice test. If a mistake

was made the procedure was repeated using the next highest concentration. If no

mistake was made we took this value as the threshold. The results of this second test

were used for the analyses presented below.


2.4. Statistical analysis

To compare thresholds of the cycling women of Group 1 tested once at different

phases of the menstrual cycle the Kruskal�/Wallis non-parametric ANOVA for

ranks was used followed by post hoc Dunn’s tests. To compare thresholds of the

cycling women of Group 2, each tested at four phases of the cycle, a non-parametric

Friedman ANOVA for repeated measures was used followed by post hoc Student�/

Newman�/Keuls tests. To compare the threshold values during ovulation of the

cycling women of Group 1 with the threshold values of the post-menopausal women

of Group 3, the pre-menstrual girls of Group 4, and the men of Group 5, non-

parametric Mann�/Whitney U tests were used. For all tests a two-tailed a value of

0.05 was taken as the level of significance.

3. Results

Fig. 1 shows the detection thresholds for amyl acetate for the 332 women ofGroup 1 tested at different phases of the menstrual cycle. Detection thresholds

differed significantly across the cycle, with a median of �/log 7.0 (range 6.0�/8.5)

during the menstrual phase, a median of �/log 7.5 during the follicular and luteal

phases (ranges 6.05/9.5 and 6.0�/9.0, respectively), and of �/log 8.5 (range 6.05/9.5)

during the ovulatory phase (Kruskal�/Wallis ANOVA; H�/18.4, df�/3, P B/0.001).

Fig. 1. Box plots of odor detection thresholds for amyl acetate for the 332 women of Group 1 tested at one

of four phases of the menstrual cycle; menstrual N�/87, follicular N�/80, ovulatory N�/21, and luteal

N�/144. Horizontal lines represent the 10th, 25th, 50th (median), 75th and 90th percentiles. *P B/0.05,

Dunn’s post hoc tests.


Thresholds recorded during the ovulatory phase were significantly lower compared

to the menstrual and luteal phases, and the follicular compared to the menstrual

phase (Dunn; P B/0.05).

As shown in Fig. 2, detection thresholds for amyl acetate also differed significantly

across the menstrual cycle for the repeatedly tested women of Group 2 (Friedman

repeated measures ANOVA; x2-value�/39.6, df�/3, PB/ 0.001). A significantly

lower median threshold of �/log 9.0 (range 8.55/9.5) was recorded during theovulatory phase compared to the threshold of �/log 8.5 recorded during the

menstrual (range 7.5�/9.0), follicular (range 8.0�/9.0) and luteal phases (range 7.5�/

8.5) (Student�/Newman�/Keuls; P B/0.05). Although the threshold scores recorded

for these women were lower than those for Group 1, the overall pattern was similar,

that is, the lowest thresholds were recorded at the time of ovulation.

As shown in Fig. 3, the median detection threshold of the women of Group 1 at

ovulation was significantly lower than that of the post-menopausal woman of Group

3 (�/log 8.5, range 6.05/9.5 vs �/log 7.0, range 6.0�/9.0, respectively; Mann�/

Whitney; U�/322, N1�/21, N2�/83, P B/0.001), and the girls of Group 4 (�/

log 8.5 vs �/log 7.0, range 6.0�/9.0; Mann�/Whitney, U�/336, N1�/21, N2�/60,

P B/0.002). Thresholds were also lower than for the men of Group 5 (�/log 8.0,

range 6.0�/9.0) although not significantly so (Mann�/Whitney, U�/1535, N1�/21,

N2�/183, P �/0.10).

4. Discussion

The present findings are consistent with those of previous studies reporting greater

olfactory sensitivity of women during the ovulatory phase of the menstrual cycle (Le

Fig. 2. Box plots of odor detection thresholds for amyl acetate of the women of Group 2 (N�/15) tested

repeatedly across one menstrual cycle. Data are presented as for Fig. 1. *P B/0.05, Student�/Newman�/

Keuls post hoc tests.


Magnen, 1952; Vierling and Rock, 1967; Mair et al., 1978; Doty et al., 1981, 1982;

Pause et al., 1994), and with reports of a late follicular or midcycle increase in

sensitivity in other sensory modalities (reviewed in Henkin, 1974; Doty et al., 1981,

1982; Brown, 1983; Doty, 1986). Thus, significantly lower detection thresholds for

amyl acetate were recorded at the time of ovulation and, at least in the large sample

of Group 1 women, the highest thresholds were recorded during menstruation.

Indeed, sensitivity at ovulation was probably even greater than represented here

given, as seen in Figs. 1 and 2, that during this phase some women could already

detect the odorant at the lowest concentration presented. Although most subjects

were tested only once and with only one substance, we consider the findings reliable

for several reasons.

The 332 women of Group 1 represent an unusually large sample compared to

earlier studies. As has been noted by previous investigators (Nghiemphu et al., 1995;

Pause et al., 1996), differences between individuals in the ability to perceive

particular odorants or in their response to the test situation may give rise to

spurious effects if sample size is small. This may partly explain the inconsistent

findings among previous reports, some of which, as mentioned in Section 1, were

based on very small samples. The large sample used here, however, made it

impracticable to test more than one odorant or to monitor individual subjects by

repeated testing across one or more cycles (cf. Henkin, 1974; Doty et al., 1981).

Fig. 3. Box plots of odor detection thresholds for amyl acetate of the cycling women of Group 1 during

the ovulatory phase (N�/21), of the post-menopausal women (N�/83), the pre-pubertal girls (N�/60),

and the men (N�/183). Data are presented as for Fig. 1. **P B/0.001, ***P B/0.0001, Mann�/Whitney U

tests.


Nevertheless, the 15 women of Group 2 tested repeatedly across the cycle in an

effort to counter the problem of single testing in Group 1, gave the same main

result*/significantly greater sensitivity around the time of ovulation. In accounting

for the lower variance and lower threshold scores of these women compared to those

of Group 1 (Figs. 1 and 2), it may be noted that the Group 2 women formed a

particularly homogeneous sample. All were military personnel living in barracks and

so were subject to the same regular daily regimen and diet and were familiar with thetest premises. The learning or practice effects noted by several authors to occur with

repeated testing (Rabin and Cain, 1986; Laska and Hudson, 1991) may also partly

explain these women’s better overall performance, although given that they were

recruited and testing started at different stages of the cycle this cannot explain their

lower thresholds during the ovulatory phase.

The specificity of the finding of higher sensitivity around the time of ovulation is

also supported by the fact that the threshold scores for the three, in several respects

very different control groups of post-menopausal women, pre-pubertal girls andyoung men were all higher than the scores obtained during ovulation for the

comparable Group 1 women who were also tested only once. Age alone would not

seem to account for the greater sensitivity of cycling women around ovulation given

their lower thresholds compared to their age peers in Group 1 tested at other phases,

or to themselves (Group 2) when tested at other phases.

And finally, although the sniff-bottle technique and the simple procedure of

ascending concentrations used here have their shortcomings such as lack of control

over the precise quantity of odorant delivered to subjects per sniff, we consider themadequate for our purpose since we were not interested in absolute sensitivity per se

and systematic errors of methodology should have affected the performance of the

groups in a similar fashion. Furthermore, sniff bottles have the advantage that

subjects can use their own natural mode of stimulus application and sniffing, and

have been repeatedly shown to generate stable and reproducible measures of

olfactory sensitivity (Laska and Hudson, 1991).

In summary, it seems clear that olfactory sensitivity varies across the menstrual

cycle, with the most consistent change represented by a decrease in threshold aroundthe time of ovulation. Nevertheless, assuming this is correct how is it to be explained?

An obvious possibility mentioned by several investigators (Le Magnen, 1952;

Schneider et al., 1958; Henkin, 1974; Good et al., 1976) is that it is due to changes in

the level of gonadal steroids, particularly estrogen. Since these hormones or their

receptors have been found in the olfactory epithelium, olfactory bulb, and other

olfactory-related brain areas in several mammalian species (Stumpf and Sar, 1982;

Vannelli and Balboni, 1982; Kratskin, 1995; Zhong et al., 2001), and we have

recently reported cytological changes in the nasal epithelium across the menstrualcycle paralleling those in the vagina (Navarrete-Palacios et al., 2003), changes in

hormone levels could conceivably influence olfactory function both peripherally and

centrally. Arguing against such a hormonal explanation, however, are findings that

women taking the contraceptive pill also show changes in olfactory threshold across

the cycle (Doty et al., 1981, 1982), that replacement therapy with estrogen has no

effect on olfactory sensitivity in post-menopausal women (Hughes et al., 2002), and


that in the present study the three hormonally different control groups did not differ

significantly in their detection thresholds for amyl acetate. Thus, although the

gonadal steroids do not exhaust the list of neurochemically active substances which

could have cycle-dependent effects on olfactory function (Doty et al., 1981; Stumpf

and Sar, 1982; Doty, 1986; Pause et al., 1996; Compagnone and Mellon, 2000), we

still lack an adequate explanation of the processes underlying these changes.

Nor is it clear what, if any, is the biological significance of such changes.Suggestions that increased olfactory sensitivity at ovulation might facilitate mate

finding or mate choice (Doty, 1974; Henkin, 1974; Benton, 1982; Grammer, 1993;

Gangestad and Thornhill, 1998; Pause et al., 1999) so far lack firm empirical

support, although reports that women can chose among the odors of men based on

small differences in the degree of match between their own and the donors’ human

leukocyte antigen alleles and in a manner consistent with enhanced immunocompe-

tence of their offspring, lends tantalizing support to this idea (Wedekind et al., 1995;

Wedekind and Furi, 1997; Jacob et al., 2002). More parsimoniously, however, itremains possible that changes in perceptual performance across the menstrual cycle

are simply a (presumably harmless) by-product of other, more fundamental aspects

of reproductive physiology. But even given this later possibility, the phenomenon

remains of interest in at least two respects*/as a factor to be taken into account in

olfactory testing, and as an indicator, as in other sensory modalities, of so far

unexplained fluctuations in the central nervous function of cycling women.

Acknowledgements

We thank Olga Alejandra Donatti for technical support, Alejandro Domınguez

for help with the figures, Carolina Rojas Castaneda for bibliographical assistance,

and the Hospital Central Militar for providing the facilities for conducting the tests.The project was supported by grants from CONACyT (25193-N) and DGAPA

(IN209999).

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Lower olfactory threshold during the ovulatory phase of the menstrual cycle

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