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Sex Aftereffects 1





Running head: FACE SEX AFTEREFFECTS

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E Sex Aftereffects in Face Perception: What You See is Not What You Get

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SS Jay Friedenberg, Megan Kwasniak, and Christine Haber

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Manhattan College

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First Author Contact Information:

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Jay Friedenberg, Ph.D.

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Associate Professor

Manhattan College

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Department of Psychology

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Riverdale, NY 10471

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Office: (718) 862-7895

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Fax: (718) 405-3249

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Sex Aftereffects 2





E-mail: Jay.Friedenberg@Manhattan.Edu

Sex Aftereffects 3





Abstract



We demonstrate a face sex aftereffect in which participants who view an

adapting face of one sex judge a subsequently appearing androgynous face as the

opposite sex. Androgynous faces were morphs made by an equal blending of a

male and female face. The aftereffect was replicated in three experiments with

different adapting durations and with both a variable and fixed inter-stimulus

interval. Attractiveness ratings provide a further validation of the phenomenon.

In experiment 3, with near equal numbers of male and female viewers, men who

perceived the morphs as female rated them more attractive than women.

Conversely, women who perceived the morphs as male rated them more

attractive than men.

Sex Aftereffects 4





Sex Aftereffects in Face Perception: What You See Is Not What You Get



In this study, we investigate a sexual aftereffect underlying face

perception. We do this by adapting viewers first to a face of determinate sex and

then have them observe a second androgynous face of indeterminate sex, what

we call a morph. Participants judged the sex and attractiveness of the morph or

of the morph and the adapting face. Morph viewing duration was always fixed,

while adapting duration and the time between presentation of the two faces was

varied across experiments. In what follows, we review the literature on sexual

dimorphism and facial attractiveness and on aftereffects and face perception. We

then describe our technique and make predictions regarding our two dependent

measures of sex judgement and perceived attractiveness.



Sexual Dimorphism and Facial Attractiveness



Mature male and female face differences contain secondary sexual

characteristics produced by hormones like testosterone and estrogen. Adult male

faces demonstrate a larger jawbone, salient cheekbones, and thinner cheeks

(Enlow, 1990). Male faces are also known to have a more protruding browridge

(Berry & Wero, 1993). This sexual dimorphism in facial shape has been linked to

perceived attractiveness. In one evolutionary account, extreme masculine or

feminine characteristics may be considered attractive to members of the opposite

sex because they advertise genetic fitness. For example, testosterone has been

shown to suppress human immune system function (Kanda, Tsuchida, &

Tamaki, 1996). Males with very masculine facial features must therefore have

strong immune systems to buffer such a handicap and may be perceived by

females as good mating partners (Folstad & Karter, 1992). Similarly, secondary

sexual characteristics in females that are estrogen dependent indicate fitness and

are considered attractive (Singh, 1993).

Available research however fails to show that masculine male faces are

consistently considered more attractive. Little & Hancock (2002) constructed

composite male faces made by averaging three, six, or 12 faces together. The

averaging technique simultaneously produced decreased ratings of masculinity

but increased ratings of attractiveness, showing a preference for feminine traits

in their male faces. Swaddle & Reierson (2002) presented a continuum of male

faces ranging from low to high masculinity. Women selected the more masculine

end of the continuum as more dominant but not most attractive. Penton-Voak &

Perrett (2000) showed a female preference for masculine faces, but only when

conception was most likely, during the follicular phase of the menstrual cycle.

They believe women have multiple motives when it comes to selecting a sexual

partner, wanting high reproductive fitness and good genes when conception is

likely, but otherwise desiring a non-dominant partner to assist in child rearing.

Sex Aftereffects 5





Other work demonstrates that women as well as men prefer feminized

faces. Rhodes, Hickford, & Jeffery (2000) found the most attractive male and

female images for Caucasian and Chinese faces were significantly feminized.

Perrett et al. (1998) found that men and women both judge feminized versions of

male faces to be most attractive. Their hypothesized explanation for this is that

feminized faces reflect socially desirable characteristics of warmth, emotionality,

honesty, and cooperativeness. These characteristics should be desirable in a

mating partner by women, but also in a friend or colleague by men.

It is interesting then, to speculate on the perceived attractiveness of

androgynous faces that are equally masculine and feminine. Such faces are the

opposite of those employed in some of the literature, since instead of magnifying

sexually dimorphic characteristics, they reduce them. If androgynous morph

faces of the sort we use here are sexually ambiguous, we would expect lowered

ratings in comparison to faces with a normal degree of masculinity or femininity.

This is because morph faces do not have clear-cut cues guiding reproductive

action. Given the finding that feminized faces are usually preferred by both

sexes, we can make another prediction. Morphs perceived as male but with

accompanying feminine characteristics, should be judged more attractive than

morphs perceived as female with accompanying masculine characteristics. In

other words, a feminized male ought to be better looking than a masculinized

female. We address this prediction in the general discussion where the results of

all three experiments are summarized. Additional predictions regarding the

perceived attractiveness of morphs are made later in the introduction.



Aftereffects and Face Perception



Aftereffects are a widespread phenomenon in visual perception. Examples

include color aftereffects between the color opponents red-green and blue-

yellow, which allows for finer discrimination between wavelengths (Abramov &

Gordon, 1994). In motion perception aftereffects exist between motions in

opposite directions. Downward motion for instance, has an opponent relation to

upward motion (Hershenson, 1993). This helps to discriminate between motions

in different directions. Aftereffects like these can be the result of opponent-

process mechanisms where receptors for opposite attributes mutually inhibit one

another. If the neural receptor for one attribute is adapted, it becomes fatigued

and temporarily looses the ability to inhibit its opponent that becomes relatively

more active.

Aftereffects have been found for more complex visual forms as well,

although it is not clear if opponent-process mechanisms underlie them. Webster

and MacLin (1999) adapted subjects to frontal views of faces that were expanded

or contracted relative to a midpoint on the nose. Test faces then appeared

distorted in the opposite direction: if the initial face was expanded, the test face

appeared contracted and vice versa. These aftereffects transferred across

Sex Aftereffects 6





different faces, but were less pronounced when the adapting and test faces were

at different orientations, i.e. the effect was reduced between faces viewed upright

and then inverted.

Visual aftereffects for other facial attributes have also been discovered.

Leopold et al. (2001) found adaptation to a specific face identity caused an

average face to take on the "opposite" identity. They had participants adapt to a

test face and then measured identification accuracy for other faces along a

trajectory of morphed faces. The trajectory consisted of a path through a multi-

dimensional space with a mean or average face in the middle, an original face on

one side of the mean, and an "anti-face" on the other side. Identification

performance was shifted toward the original face after adaptation to the anti-

face, in comparison to adaptation to a non-matching face on another trajectory or

to a no adaptation condition. In addition Kaping et al. (2002) reports that

adaptation to a particular emotional expression (happy or sad), race (Caucasian

or Japanese), or sex (male or female) could cause a prior neutral image to acquire

the opposite characteristics.

Rhodes et al. (2003) demonstrated that contrast effects occur also for

judgments of averageness and attractiveness. They adapted viewers to faces that

were expanded or contracted relative to an average face. The participant's

consideration of what faces were average and attractive shifted in the direction of

the adaptation. These results suggest that adaptation renormalizes the face space

or dimension of facial features so that what is considered average or "normal" is

now centered closer to the adapting stimulus. They hypothesize that this type of

mechanism may have evolved to allow individuals to adjust to local population

characteristics, since non-migratory communities may be best suited to resident

environmental conditions (Rhodes et al., 2003). Alternatively, Rhodes mentions

that this sort of mechanism may be more general, reflecting the extraction of

prototypes useful for recognizing and categorizing objects (Rosch, 1973).

As Kaping et al (2002) have shown there are aftereffects for facial sexual

characteristics. The sex of faces is an extremely consequential feature. It has

specific functional implications for social interaction. For instance, knowing

someone's sex informs us whether this person poses a threat or may be a

potential reproductive partner. Cases of sexual ambiguity demonstrate just how

important this is. If we encounter an androgynous person whose sex cannot be

determined, we are at a loss as to how to behave towards them.

An aftereffect for the dimension of face sexuality has interesting

implications. Adapting to male faces would make them the new average.

Subsequently, an androgynous face with mixed features would now be judged

female, since the average has been moved toward the male end of the

continuum. Conversely, adaptation to female faces would shift the average

toward the feminine side causing androgynous faces to be perceived as male.

The consequence of this is a perceptual adaptation to the local distribution of

sexual characteristics. Time spent in a predominately male-looking social group

Sex Aftereffects 7





would predispose an individual to perceive a slightly feminine face as female.

Time spent among a female-looking group would additionally cause faces with

small amounts of masculinity to appear male. From a functional standpoint, this

adaptation would facilitate appropriate social interaction within the group.

In this study, we predict the presence of such a face sex aftereffect. We

demonstrate that continued exposure to a face of one sex produces increased

perceptual salience for the opposite sex. Continued exposure to a male face for

example produces increased salience of female face attributes. Conversely,

adaptation to a female face results in increased perceptual salience for male facial

characteristics.



Technique and Predictions



We test this aftereffect hypothesis in the current study using an adaptation

technique. Adapting faces, which are unaltered individual male or female faces,

are presented to observers for a given duration. Participants are then shown

androgynous faces. These androgynous faces contain the face previously viewed

combined with another face of the opposite sex never seen in the experiment. We

call these sexually ambiguous faces morphs, since they are constructed by an

equally weighted average of a single male and female face. If there is a face sex

aftereffect, exposure to a male adapting face should increase saliency of female

faces and vice versa. This can be measured in the observer's subsequent

judgement of the morph. If they are exposed to male adapting faces, they ought

to perceive the morphs as more female. Should they view female adapting faces,

they instead ought to see the morph as male.

In this investigation we ask observers to also rate the perceived

attractiveness, both of the adapting faces and the morphs. If there is no

aftereffect, then morph faces, with indeterminate sexual characteristics, should

be rated less attractive than unaltered adapting faces. If there is a face sex

aftereffect, then morphs will have a sexual identity and attractiveness ratings to

them should vary depending on sex of the observer. We expect that morphs

perceived as male will be judged more attractive by females and vice versa.

Finally, as discussed above we predict that morphs considered as male will be

judged better looking by both sexes.



Experiment 1



Method



Participants.



A total of 56 students from Manhattan College and the College of Mount

Saint Vincent participated. There were 16 males and 40 females from three

Sex Aftereffects 8





undergraduate classes. Average age was 19 years. All received extra credit for

their participation and had vision that was normal or corrected to normal.



Apparatus.



Adapting faces were obtained from a University of Michigan Internet face

database, now defunct. The faces were of undergraduate engineering students.

Those used in the experiment were selected on the following criteria: a full

frontal view, no obvious blemishes, no glasses or facial hair, and the absence of

strong shadows or reflections. The images were in color and cropped below the

chin and above the hairline. Facial expression varied slightly.

There were a total of 56 unique adapting faces. An adapting face was

unaltered, it did not undergo any stimulus manipulation. Adapting faces were

presented to observers but also used to generate the morphs. An attempt was

made to balance adapting faces with respect to ethnicity. This resulted in 14

Asian, 20 Black, and 22 Caucasian faces. Given our sample, there were an

insufficient number of adapting faces to create Hispanic morphs, so Hispanic

faces were not used.

We created morph faces by equally combining a single male and a single

female adapting face. This yielded 28 total morphs, seven Asians, 10 Blacks, and

11 Caucasians. The two single adapting faces were superimposed and blended to

create a 50% composite image using the Morpher 2.0 program for the Mac

(Fujimiya, 1997). In Morpher, the user selects nodes, which are corresponding

points on the two input images at high regions of contrast. These nodes on each

image are then connected with lines called boundaries. Morpher uses the

corresponding boundary information on each image to create a composite image,

which is weighted pixel average.

All faces, adapting stimuli and morphs, were presented in color on an

Apple iBook laptop computer. Face dimensions were 4.0 cm in width and 4.5 cm

in height. Participants made judgements at an average viewing distance of 45 cm.

Each face was centered in the middle of the 35.5 cm (14 inch) diagonal length

screen. A 50% homogenous gray background filled the remainder of the viewing

region.



Procedure.



Data collection from the three classes took place in one classroom. Each

class was run in one or two sessions of approximately 15 students each.

Participants first read and signed a consent form then read a detailed set of

written instructions. After this, the research assistant explained the instructions

to them verbally and answered any questions they had about the procedure.

The three conditions in the study were Male Adapt, with male adapt faces

followed by morphs, Female Adapt, with female adapts preceding morphs, and

Sex Aftereffects 9





the No Adapt condition, with morph faces only. The students were randomly

assigned to one of these three conditions, but did not know which condition they

were in. Instructions for each condition were the same, to simply view each face

that appeared and evaluate it, first with respect to sex, then with respect to its

perceived attractiveness.

Each participant viewed the experiment on his or her own monitor. In the

No Adapt condition, morphs appeared for seven seconds. A response screen

then appeared prompting the participant to push the "M" key if they thought it

was a male face and the "F" key if it was perceived as female. A second response

screen then appeared prompting the participant to enter a number between "1"

and "7" to indicate the perceived attractiveness of the face, with higher numbers

indicating greater perceived attractiveness. Both response screens were response

terminated and remained up for as long as it took an individual to decide. The

time to make each decision was recorded. Order of faces was randomized.

The two adapt conditions contained twice as many trials as the No Adapt

conditions, because participants first viewed and evaluated an adapting face and

then its corresponding morph. An adapting face appeared for seven seconds,

followed by the sex and attractiveness rating screens. Then in the next trial the

morph face appeared also for seven seconds and followed by the two rating

screens. The adapt face that preceded a morph was always one of the two

adapting faces that were used to construct it. In the male adapt condition it was

the male face. In the female adapt condition it was the female face. The order of

these adapt-morph trial pairs was randomized.



Results and Discussion



The primary independent variable was Adapt Type, with three levels.

These were Male Adapt, Female Adapt, and the No Adapt control. Adapt Type

was a between-subjects variable. There were two major dependent variables. The

first was sex judgement. This was the observer's perception of whether the faces

were male or female. The second were attractiveness ratings. This was the

perceived attractiveness of the faces based on the seven-point rating scale. The

results of each dependent variable are summarized in separate sections below.

The data was first screened for outliers. All responses greater than seven

seconds, which constituted less than one percent of the data, were excluded from

subsequent analyses.



Sex Judgements.



Chi-square tests were performed separately for morph and adapt faces.

This was done because all three conditions contain morphs but only the male and

female adapt conditions have adapting faces. There was a significant effect of

Adapt Type on the perceived sex of morph faces 2 (2, N = 1,549) = 72.43, p <

Sex Aftereffects 10





0.01. Observers judged the morphs in the No Adapt condition ambiguous with

respect to sex, with a 55% - 45% male - female split. These data demonstrate that

the morph faces were perceived as androgynous, with perhaps a slight male bias.

The results also indicate the presence of a face sex aftereffect. Sixty percent of the

morphs were judged female when preceded by a male adapt, while 67.3% of

them were perceived male when they followed a female adapt. These data are

consistent with our hypothesis that exposure to the adapting faces may fatigue a

sex-specific face representation, allowing the opposite sex characteristics of the

morph to become salient.

As expected, there was a highly significant effect of Adapt Type on sex

judgements for the adapt faces, 2 (1, N = 925) = 812.81, p < 0.01. Adapting faces

were very unambiguous with respect to sex with 97.8% responding male in Male

Adapt and 95.8% responding female for Female Adapt. This demonstrates that

our unaltered adapting stimuli were categorically perceived as distinctly male or

female. Table 1 shows the percentage of male and female responses in the Adapt

Type conditions for both types of face.



Attractiveness Ratings.



As before, analyses were performed separately for adapts and adapting

faces. A one-way ANOVA showed there was a significant effect of Adapt Type

on perceived attractiveness for morphs F(2, 53) = 44.23, p < 0.01. Morphs in the

Female Adapt group were considered most attractive (M = 3.41, SE = 0.14).

Morphs in the Male Adapt condition were rated least attractive (M = 2.63, SE =

0.13). Morphs in the No Adapt control fell in between (M = 3.14, SE = 0.08) All

pair-wise comparisons between these means were significant by Fisher's

Protected Least Significant Difference (PLSD) at an alpha level of 0.05. Notice

that morph faces alone in the No Adapt condition were not rated as less

attractive than all of the adapt faces.

An explanation for the Adapt Type variable results emerges when the

data are broken down by sex of the participant. Women judged morphs in the

Female Adapt group as more attractive (M = 3.59, SE = 0.07) than men (M = 3.04,

SE = 0.12). Because the female adapts masculinized the morphs, heterosexual

women should see them as more attractive. Since there are more than twice as

many women as men in our sample, female judgements are probably exerting a

stronger influence on the data, raising the overall mean in this condition. In the

Male Adapt condition, feminization of the morphs would produce an opposite

effect. Here we would expect men to rate the morphs higher. In fact they don't,

but the mean difference between men (M = 2.49, SE = 0.10) and women (M =

2.67, SE = 0.07) is much smaller. This may be due to the limited number of male

participants. In experiments 2 and 3, with a more equal number of observers

from either sex, differences in responding to these conditions disappears.

Sex Aftereffects 11





The separate analysis for adapting faces showed greater perceived

attractiveness in the Female Adapt condition (M = 3.35, SE = 0.06) than for Male

Adapt (M = 2.85, SE = 0.06). This was a significant difference t(32) = 28.54, p <

0.01. When subsequently broken down by sex of participant, it turns out this

larger mean comes from female (M = 3.55, SE = 0.07) rather than male (M = 2.89,

SE = 0.12) observers.

Experiment 2



Aftereffect phenomena require exposure to an adapting stimulus. But the

actual amount of adaptation needed to elicit aftereffects varies. In some cases, it

is only a few seconds. Webster and MacLin (1999) obtained figural aftereffects for

faces with just a few seconds of exposure time. So did Rhodes et al. (2003). In

others, like chromatic adaptation, exposure time may run 45 seconds or higher

(Uchikawa, K, Uchikawa, H., & Boynton, R. M., 1989).

In experiment 1 we obtained aftereffects with as little as seven seconds of

adaptation time. In the next experiment we employ adaptation times both below

and above seven seconds to see if the strength of the aftereffect changes. In

particular, we wish to see if there is a minimum exposure time necessary to elicit

the phenomenon and whether it will increase with an increase in adapting

stimulus view time. Hershenson (1993) found that motion aftereffects are more

powerful the longer one looks at the inducing stimulus. If the face aftereffect is

governed by the same mechanisms as in these well-known aftereffects, we could

expect greater adaptation to produce a stronger face sex aftereffect.

In experiment 2 we employed a short adaptation duration of two seconds,

five seconds below that used in experiment 1 and a longer adapting duration of

12 seconds, five seconds above that used previously. It is unclear what to predict

for the 2-second condition. If this viewing time is too short it may be insufficient

to elicit any type of adaptation and there will be no effect. A 12-second exposure

though, should allow for greater adaptation and so produce an aftereffect

stronger than that seen in experiment 1.



Method



Participants.



A different set of 56 Manhattan College and College of Mount Saint

Vincent undergraduate students participated for extra class credit. There were 27

males and 29 females from three separate psychology courses. Average age was

19.6 years. Vision was normal or corrected to normal.



Procedure.

Sex Aftereffects 12





Unless otherwise stated, the stimuli and procedure used here were

identical to that of experiment 1. There were four between-subject conditions:

Male Adapt 2-seconds, Male Adapt 12-seconds, Female Adapt 2-seconds, and

Female Adapt 12-seconds. These times were deliberately set to be symmetrically

five seconds below and above the experiment 1 duration. Viewing duration of

the morph faces that followed was seven seconds, the same as in the previous

study. Response screens for sex judgement and attractiveness ratings were

response terminated as before. In the male adapt conditions, participants saw

only the male adapt faces. In the female adapt conditions they saw only the

female adapt faces. Order of the adapt-morph trial pairs was randomized.

Participants were randomly assigned to one of the four conditions and were

naive with respect to each condition. Running time for the 2-second versions was

20 minutes. For the 12-second versions, it was 25 minutes.



Results



There were two primary independent variables. These were Adapt Sex

with two levels, Male Adapt and Female Adapt. This was crossed with Adapt

Duration, also with two levels, 2-seconds and 12-seconds. Dependent measures

were the same as in experiment 1. Responses greater than seven seconds were

considered outliers and removed from all analyses. They made up less than one

percent of the data.



Sex Judgements.



The morph data were examined first. There was a significant effect of the

Adapt Sex variable 2 (3, N = 1,607) = 44.75, p < 0.01. As expected, the

distribution of male and female responses to morphs were about equal in the

short viewing conditions. For Male Adapt 2-seconds and Female Adapt 2-

seconds, percent responding to either sex was very close to an even 50 - 50 split.

In the longer viewing conditions, there was again evidence of a face sex

aftereffect. For the Male Adapt 12-second condition, 63.5% of the morphs were

perceived as female. In the Female Adapt 12-second condition, 58.5% of the

morphs were seen as male. Table 2 shows the percentage of male and female

responses for the Adapt Sex and Adapt Duration variables for both types of face.

The Adapt Sex variable had a significant effect on responding to adapting

faces faces as well, 2 (3, N = 1,604) = 1,409.63, p < 0.01. As expected, the adapts

were unambiguous. There was near unanimity in perception of male faces as

male and female faces as female. There was no discernible difference in

responding to these faces as a function of duration, i.e., participants were not

substantially more accurate at longer durations.

These data require some discussion. Our first major prediction, that a

short duration would not elicit the face aftereffect, was fulfilled. Responding in

Sex Aftereffects 13





these cases was identical to a 50 - 50 split. This is consistent with the assumption

than an internal face representation must be adapted to produce the aftereffect.

Two seconds of viewing time is apparently too little time to do so. Our second

prediction, that the aftereffect should be stronger for longer adapting durations,

was not entirely fulfilled. In experiment 1, the Male Adapt condition produced a

60% female response to morphs. The current data show a slight increase to

63.5%. In experiment 1, there were 67.3% male responses to morphs in the

Female Adapt condition. In the current study with longer durations this drops to

58.5%. If one collapses across Adapt Sex, there was an overall greater sex

aftereffect in the first study where duration was seven seconds.

Why is this the case? Twelve seconds is still a fairly short adaptation time

compared to other perceptual aftereffect phenomena. These results suggest a fast

adapting system for faces, one that peaks somewhere between the two values

tested here, but which may decline shortly thereafter. A fast adapting system

may match the rapid pace of social discourse. Important linguistic ideas are often

communicated in just a few seconds. Sex appropriate discourse may thus also

require rapid sexual discrimination. We cannot distinguish here whether this

result is due to a fast-acting adaptation or simply an artifact of the way in which

the visual system works. The exact time-course of this effect warrants additional

investigation. We study it further in the next experiment.



Attractiveness Ratings.



First of all, we note that morph faces overall were not rated significantly

less attractive than adapts, again suggesting that morph faces are not entirely

sexually indeterminate, as would be expected if there were a face sex aftereffect

at work. A (2 x 2) factorial ANOVA with Adapt Sex and Adapt Duration as

factors was performed next for the attractiveness ratings. This was done for

morph faces first. There was no effect of Adapt Sex. Ratings were not

significantly greater for female adapts. There was a significant effect of Adapt

Duration F(1, 44) = 3.57, p < 0.05, with morphs in the longer 12-second duration

judged more attractive (M = 2.91, SE = 0.04) than those in the 2-second duration

(M = 2.84, SE = 0.05). The Adapt Sex by Adapt Duration interaction was not

significant.

This same analysis was next applied to the adapt faces. Adapt Sex and the

Adapt Sex by Adapt Duration interaction were not significant. There was again a

main effect of Adapt Duration F(1,44) = 4.56, p < 0.05. Adapts viewed for 12

seconds were judged more attractive (M = 2.99, SE = 0.04) than those viewed for

two seconds (M = 2.84, SE = 0.05).

Why are the adapting faces more attractive when viewed longer?

According to the mere exposure effect, liking for novel stimuli, including human

faces, increases with repeated exposure (Moreland & Zajonc, 1982). Longer

viewing times may increase familiarity and therefore perceived attractiveness in

Sex Aftereffects 14





these faces. But how can we explain this same result for morphs? They were

viewed for the same time in both the short and long conditions. It is possible that

ratings for adapts may generalize to morphs. When an adapting face is judged

better looking, it increases the probability that the morph, which resembles it,

will also be considered better looking.



Experiment 3



In the first two experiments, observers judged both faces. This was

necessary since it allowed us to compare the relationship between them.

However, judgements to the first face produced a variable time interval between

it and the second face. Across all viewers in experiment 1 average decision time

for the two response screens was a little over two seconds. This variability makes

it difficult to ascertain a pure effect of adapting duration. Although the first face

appears for a fixed time, its subsequent influence may fade, especially at a longer

inter-stimulus interval (ISI).

To remedy this situation, we conduct a third study where the duration

between adapt offset and morph onset is fixed. In fact, we eliminate any ISI

altogether with the morph appearing immediately after the adapt. We predict the

elimination of the ISI should produce stronger aftereffects. We employ the same

adapting durations as in the previous studies: two, seven and 12 seconds.

Experiment 3 thus serves as a controlled replication of the first two experiments.

In addition, an attempt was made this time to deliberately equate the

number of male and female participants in each condition, allowing us to

examine the interaction of this subject variable. The inclusion of participant sex

in the analysis also gives us a means to validate the opposite sex effect. If male

adapts induce perception of a female morph, then heterosexual male viewers

should rate these morphs higher in attractiveness. Conversely, female adapts

ought to induce higher attractiveness ratings of morphs by heterosexual female

viewers if they are perceived as male.



Method



Participants.



A total of 82 undergraduates from Manhattan College and the College of

Mount Saint Vincent participated for extra class credit. There were 36 males and

46 females. Average age was 19.4 years. Vision was normal or corrected to

normal.



Procedure.

Sex Aftereffects 15





Adapts and morphs were the same as those used in the first two studies.

Participants were instructed to view both faces in a trial but to evaluate only the

second face of each pair. Adapt duration varied depending on condition. Morphs

immediately superceded adapting faces. Morph duration remained at seven

seconds. The two response screens followed presentation of the morphs.

Six conditions resulted from the crossing of Adapt Duration with three

levels (2-seconds, 7-seconds, and 12-seconds) and Adapt Sex with two levels

(Male Adapt and Female Adapt). Male and female observers were separately and

randomly assigned to these six conditions. On average there were seven

members of each sex in each of these six groups. All other procedural aspects

matched those used earlier.



Results and Discussion



Adapt Duration and Adapt Sex were the independent variables. Sex

judgement, attractiveness ratings, and RT were dependents and are reported in

each section below. Participant Sex was a subject variable. Outliers were screened

following the same criteria as before. The data in this experiment are responses

to morph faces only.

Sex Judgements.



Chi square analyses for sex judgement were performed separately for

Male Adapt and Female Adapt. There was a significant effect of Adapt Duration

for female faces, X2 (2, N = 1,204) = 10.53, p < 0.01. The effect for male faces was

marginally significant, X2 (2, N = 1,118) = 3.82, p = 0.14. For the 2-second duration

for male and female adapts, responding to morphs approximated a 50 - 50 split.

For the longer duration of 7-seconds and 12-seconds, there is a clear aftereffect at

work in each case approximating a two-thirds split. Table 3 shows percent

responding to male and female adapts for each level of Adapt Duration.

In comparing these values to those obtained with a variable ISI, one sees

little difference. In experiment 2, the 2-second duration with a variable ISI

produced responding indistinguishable from a 50 - 50 split, with deviations away

from this of no more than two percent. In the current study with a fixed and

immediate ISI, the deviations from a perfect 50 - 50 split are a bit larger,

bordering on five percent. These numbers however do not differ substantially

from the No Adapt control data in experiment 1, where there was a 55 - 45 male -

female split. We conclude therefore that a 2-second adapt duration, even with a

zero ISI is insufficient to elicit the effect.

There is also little difference in the judgement data when comparing the

longer adapting durations. The 7-second conditions for male and female adapts

in experiment 1 are identical to those obtained here. For the 12-second duration,

the differences in responding are also negligible, within a few percentage points.

One can conclude from these results that the variable ISIs in the first two

Sex Aftereffects 16





experiments were of such a short nature that they did not allow for any

meaningful loss of the effect.

Additional chi square analyses were next performed for Adapt Duration

and Participant Sex. There were no major differences in the pattern of statistical

significance or of responding by male or female observers. This suggests that the

mechanism underlying the aftereffect is the same in both sexes. If the putative

reason for such a mechanism is the ability to make fine-tuned discriminations

between members of each sex so as to functionally adjust one's social behavior,

then there is no reason to expect any such difference. Members of both sexes

have an equally important need to recognize and identify the sex to which a

given individual belongs. The primary reasons for this identification are the

same in men and women, namely threat and mate assessment.

To summarize, the data from this experiment replicate quite well the

equivalent conditions from experiments 1 and 2. The pattern of response for

judging the sex of morph faces is the same in all cases. There is an absence of any

aftereffect at short 2-second intervals, an apparent asymptote at 7-seconds, and a

leveling off of the effect at adapt durations greater than this. In addition, there is

no discernible difference in the face sex aftereffect for male and female observers.



Attractiveness Ratings.



A factorial ANOVA with Adapt Sex and Participant Sex as factors was

calculated in order to examine any interaction. There was no main effect of

Adapt Sex. There was a significant main effect of Participant Sex, F(1, 54) = 5.70,

p < 0.05. Female observers judged the morphs more attractive (M = 3.02, SE =

0.04) than males (M = 2.87, SE = 0.04). The interaction between Adapt Sex and

Participant Sex was also significant, F(1, 54) = 54.36, p < 0.01.

The shape of the interaction between sex of adapt and participant is in the

form of a cross over. In the Female Adapt condition where morphs are perceived

as male, females rate them more attractive (M = 3.24, SE = 0.05) than males (M =

2.70, SE = 0.06). In the Male Adapt condition where morphs are perceived as

female, males rate them higher (M = 3.05, SE = 0.06) than females (M = 2.75, SE =

0.05). This finding nicely validates the aftereffect. It shows that in a primarily

heterosexual subject pool, morphs biased toward male are considered more

attractive by females. Conversely, morphs biased toward female are rated more

attractive by males. This validation is only appropriate here in this experiment,

where the number of observers of each sex were equated.



General Discussion



The Face Sex Aftereffect

Sex Aftereffects 17





In the introduction, we hypothesized the existence of a face sex aftereffect,

where exposure to a face of one sex would make a subsequently appearing

androgynous morph appear to be the opposite sex. The two instances of this are

viewing a male adapting face and perceiving the morph as female and viewing a

female adapting face and perceiving the morph as male. These specific

predictions were confirmed in three separate experiments under a variety of

different conditions, showing the phenomenon is fairly robust.

We also manipulated duration of the adapting face, expecting that longer

durations would produce a stronger aftereffect. This prediction was partly

confirmed. The results of experiments 1 and 2 show no aftereffect at a short 2-

second duration, a strong effect at a moderate 7-second duration, but no

continued increase in the magnitude of the effect at a longer 12-second duration.

We conclude that 2-seconds is insufficient time to produce the effect but that it

may occur quite quickly thereafter, reaching a peak only a few seconds later. The

speed of this adaptation, as discussed earlier, may be related to the rapid

requirement for sexual identification in social discourse. Future research might

investigate aftereffect strength as a function of shorter intervals in the range we

used here as well as at longer durations beyond 12-seconds.

In the first two experiments, observers judged adapting faces and morphs.

This introduced a variable ISI between faces that could have mitigated the

aftereffect. To see if this was the case we conducted a third experiment with a

fixed and immediate ISI. The magnitude of the aftereffect remained unchanged,

showing that a spacing of several seconds in between faces is not enough to

diminish the phenomenon. Experiments 1 and 2 did not control for sex of the

observer. So in experiment 3 we tested equal numbers of males and females,

finding no difference in the aftereffect for participant sex. We conclude based on

these results that the aftereffect is the same in both sexes because its functional

significance for both sexes is the same.

We reiterate the reason for such a mechanism as stated in the introduction.

Appropriate social interaction requires unambiguous sexual identification.

Adaptation to a local population of sexual face characteristics normalizes an

individual's responses to that group. This reduces ambiguity and promotes

adaptive sex-specific social behavior. Our data suggest that time spent among

males for example will renormalize an internal face sex dimension so that the

mean is now shifted closer to males. This makes subsequently appearing faces

with smaller amounts of femininity appear more female. Men under these

conditions would be more likely to mate with these females, a response

promoting reproduction and group survival. Women under these conditions

would also perceive these faces as more female and react in a socially

appropriate manner, perhaps through increased cooperative interaction that

would also promote group survival. If this explanation is correct, then the face

normalization process we report here may be the result of evolutionary selection

pressures.

Sex Aftereffects 18





What kind of mechanism underlies this phenomenon? Eifuku et. al. (2004)

have found that the anterior inferior temporal gyrus (ITG) in macaque monkeys

is involved in judgments of facial identity in conjunction with the superior

temporal sulcus (STS). In humans the STS is implicated in processing of facial

movement and expression both of which are social cues (Allison, Puce, &

McCarthy, 2000). It is possible neurons in these or other regions code for sexual

identity of faces and that these cells might have opponent relations. If this were

the case, activation in the cell population coding for one sex might suppress

activity in neurons coding for the opposite sex. This could mediate the effect we

have discovered here.

We want to emphasize that the above explanation is speculative. This

investigation does not address the neural substrate of the phenomenon. It is still

not known if the aftereffect is mitigated by mutual inhibition between neural face

detectors or is the result of some other process. This study only demonstrates the

existence of the aftereffect and does not attempt to explicate the brain

mechanisms that give rise to it. There is in fact some doubt about the exact nature

of any such mechanism. The effect seems to be mediated by high-level neurons

that are not tied to retinotopic input, since Rhodes et al. (2003) obtained

aftereffects with a 90 change in orientation between test and adapting faces.

Another question is whether the phenomenon is even specific to faces at all. The

face adaptation effects obtained here and elsewhere may be the result of a

general-purpose perceptual process that mediates a wide variety of different

stimuli.



Attractiveness Ratings



We made several speculations with regard to attractiveness in the

introduction. First, we stated that morphs might not be rated less attractive than

the adapting faces if they were perceived as male or female. The data confirm

this. In the first two experiments, there was no significant difference in ratings

between adapts and morphs. This is what one would expect if the morphs were

not perceived as androgynous because of the influence of the preceding faces.

Second, we used the dependent measure of attractiveness to validate the

face sex aftereffect. Assuming a primarily heterosexual subject pool, we

predicted that males should judge morphs higher in the Male Adapt condition

where the morphs are perceived as female. Alternatively, females should judge

morphs higher in the Female Adapt condition that biases morphs as male.

Overall though, there should be no difference in ratings for the Adapt Sex

variable across both sexes, since opposing male and female preferences should

cancel each other out. These predictions were confirmed in experiments 2 and 3

where approximately equal numbers of male and female viewers were tested.

They were to some extent also corroborated in experiment 1. In that study,

Sex Aftereffects 19





ratings were higher in the Female Adapt condition that seems to have been due

to the large number of female viewers.

In the introduction, we predicted that morphs perceived as male would be

judged more attractive than morphs perceived as female. The literature shows

observers consider feminized faces more attractive, perhaps because these

features signal cooperativeness and other socially desirable characteristics. Was

this the case in our study? In experiments 1 and 3, there was a slight tendency to

judge morphs perceived as male more attractive. In experiment 2, this trend was

reversed. In all cases, the mean differences were not significant. We conclude

then that morphs of the sort we use here are not considered more attractive when

judged male.



Future Prospects



There are a few limitations in the current study. The aftereffect

demonstrated here is specific to the faces we used. In our stimuli, the adapting

face constituted half of the morph. It remains to be seen if the aftereffect is more

general, linked to a generic representation of maleness or femaleness and not to

any other particular facial characteristics. Recall that the figural aftereffects

obtained by Webster and MacLin (1999) transferred across different faces. These

results imply that aftereffects for sexual identity may also be general. In any

event, this hypothesis can be tested by constructing morphs from male and

female faces that are not used as adapting stimuli in the same experiment. If the

aftereffect is undiminished then the neural representations being adapted are not

tied to any individual configuration of facial features and instead code for an

abstract form of sexual identity. If the aftereffect is diminished, one could then

vary the similarity of the adapting face to its morph counterpart to see which face

attributes the effect may be linked. This would be an interesting investigation, as

it addresses the question of whether sexual identity is a separable or integral face

dimension (Garner, 1974).

We should mention here that morphs have limited ecological validity,

since faces of this sort are so rarely encountered in everyday life. It is of course

possible to evaluate the aftereffect without using morphs at all. One could do this

by presenting a male or female adapt and then following it with a normal

unaltered male or female face. Participants instead of judging sex would rate the

masculinity or femininity of the second face. Our data predict that male adapting

faces would feminize the ensuing faces while female adapts would masculinize

them.

Sex Aftereffects 20





References



Abramov, I., & Gordon, J. (1994). Color appearance: On seeing red, or



yellow, or green, or blue. Annual Review of Psychology, 45, 451-485.



Allison, T., Puce, A., & McCarthy, G. (2000). Social perception from visual cues:



Role of the STS region. Trends in Cognitive Sciences, 4, 267-278.



Berry, D. S., & Wero, J. L. F. (1993). Accuracy in face perception: A view



from ecological psychology. Journal of Personality, 61, 497-520.



Eifuku, S., De Souza, W. C., Tamura, R., Nishijo, H., & Ono, T. (2004). Neuronal



correlates of face identification in the monkey anterior temporal cortical



areas. Journal of Neurophysiology, 91, 358-371.



Enlow, D. H. (1990). Facial growth (3rd ed.). Philadelphia: Harcourt



Brace Jovanovich.



Folstad, I., & Karter, A. J. (1992). Parasites, bright males and the



immunocompetence handicap. American Naturalist, 139, 603-622.



Fujimiya, M. (1997). Morpher for Macintosh. Retrieved August 23, 2001,



from http://www.asahi-net.or.jp/~FX6M-FJMY/mop00e.html.



Garner, W. (1974). The processing of information and structure. New Jersey:



Lawrence Erlbaum.



Hershenson, M. (1993). Linear and rotational motion aftereffects as a



function of inspection duration. Vision Research, 33(14), 1913-1919.



Kanda, N., Tsuchida, T., & Tamaki, K. (1996). Testosterone inhibits



immunoglobulin production by human peripheral blood

Sex Aftereffects 21







mononuclear cells. Clinical and Experimental Immunology, 106, 410-



415.



Kaping, D., Bilson, A.C., & Webster, M. A. (2002, November). Adaptation and



categorical judgments of faces. Poster presented at the Object Perception



and Memory Conference, Kansas City, MO.



Leopold, D. A., O'Toole, A.J., Vetter, T., & Blanz, V. (2001). Prototype-referenced



shape encoding revealed by high-level aftereffects. Nature Neuroscience,



4, 89-94.



Little, A. C., & Hancock, P. J. B. (2002). The role of masculinity and



distinctiveness in judgments of human male facial attractiveness.



British Journal of Pychology, 93, 451-464.



Moreland, R. L., & Zajonc, R. B. (1982). Exposure effects in person



perception: Familiarity, similarity, and attraction. Journal of



Experimental Social Psychology, 18, 395-415.



Penton-Voak, I., Perrett, D. (2000). Female preference for male faces



changes cyclically-further evidence. Evolution and Human



Behaviour, 21, 39-48.



Perrett, D. I., Lee, K. J., Penton-Voak, I. S, Rowland, D. R., Yoshikawa, S.,



Burt, D. M., Henzi, S. P., Castles, D. L., & Akamatsu, S. (1998).



Effects of sexual dimorphism on facial attractiveness. Nature, 394,



884-887.

Sex Aftereffects 22





Rhodes, G., Hickford, C., & Jeffery, L. (2000). Sex-typicality and



attractiveness: Are supermale and superfemale faces



superattractive? British Journal of Psychology, 91, 125-140.



Rhodes, G., Jeffery, L., Watson, T. L., Clifford, C. W. G., & Nakayama, K. (2003).



Fitting the mind to the world: Face adaptation and attractiveness



aftereffects. Psychological Science, 14(6), 558-566.



Russell, J. A., & Carroll, J. M. (1999). On the bipolarity of positive and



negative affect. Psychological Bulletin, 125(1), 3-30.



Singh, D. (1993). Body shape and women's attractiveness-the critical role



of waist-to-hip ratio. Human Nature, 4, 297-321.



Swaddle, J. P., & Reierson, G. W. (2002). Testosterone increases perceived



dominance but not attractiveness in human males. Proceedings of



the Royal Society of London. Series B., 269, 2285-2289.



Uchikawa, K., Uchikawa, H., & Boynton, R. M. (1989). Partial color



constancy of isolated surface colors examined by a color-naming



method. Perception, 18, 83-91.



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perception of faces. Psychonomic Bulletin & Review, 6, 647-653.

Sex Aftereffects 23





Author Note



Jay Friedenberg, Megan Kwasniak, and Christine Haber, Department of

Psychology, Manhattan College.

Preliminary results of the first two experiments were presented at the 43rd

Annual Meeting of the Psychonomic Society, November 21 – November 24, 2002,

in Kansas City, Missouri.

Correspondence concerning this article should be addressed to Jay

Friedenberg, Department of Psychology, Manhattan College, Manhattan College

Parkway, Riverdale, New York 10471. Electronic mail may be sent via Internet to

Jay.Friedenberg@Manhattan.Edu

Sex Aftereffects 24





Table 1



Percent responding to morph and adapting faces in experiment 1 for type of

adapt

______________________________________________________________________



Adapt Type

____________



Response Female Adapt Male Adapt No Adapt

_____________________________________________________________________



Morph Faces



Male 67.4 39.9 55.2



Female 32.6 60.1 44.8



Adapt Faces



Male 4.2 97.9 N.A.



Female 95.8 2.1 N.A.

_____________________________________________________________________

Sex Aftereffects 25





Table 2



Percent responding to morph and adapting faces in experiment 2 as a function of

adapt sex and duration

_____________________________________________________________________



Adapt Sex



Female Adapt Male Adapt

______________ ____________



Adapt Duration 2 12 2 12

(seconds)

_____________________________________________________________________



Morph Faces



Male 48.2 58.5 50.5 36.6



Female 51.8 42.2 49.5 63.4



Adapt Faces



Male 2.7 4.7 97.0 97.9



Female 97.3 95.3 3.0 2.1

_____________________________________________________________________

Sex Aftereffects 26





Table 3



Percent responding to morph faces in experiment 3 as a function of adapt sex and

duration

______________________________________________________________________



Adapt Sex



Female Adapt Male Adapt

_____________ ____________



Adapt Duration 2 7 12 2 7 12

(seconds)

_____________________________________________________________________



Response



Male 56.5 66.7 65.4 45.2 39.5 38.7



Female 43.5 33.3 34.6 54.8 60.5 61.3

_____________________________________________________________________

Sex Aftereffects 27

Sex Aftereffects 28

Sex Aftereffects 29


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