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C. Bushdid et al.
Science 343, 1370 (2014);
DOI: 10.1126/science.1249168
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REPORTS
1
Laboratory of Neurogenetics and Behavior, The Rockefeller
University, 1230 York Avenue, Box 63, New York, NY 10065,
USA. 2Laboratory of Mathematical Physics, The Rockefeller
University, 1230 York Avenue, Box 212, New York, NY 10065,
USA. 3Howard Hughes Medical Institute, 1230 York Avenue,
Post Office Box 63, New York, NY 10065, USA.
90% overlap
60% overlap
30% overlap
No overlap
1.34x1017 possible
mixture pairs
1.83x1023 possible
mixture pairs
7.18x1026 possible
mixture pairs
1.10x1028 possible
mixture pairs
95% overlap
75% overlap
50% overlap
25% overlap
No overlap
1.29x1026 possible
mixture pairs
1.03x1035 possible
mixture pairs
4.28x1042 possible
mixture pairs
8.12x1044 possible
mixture pairs
1.76x1044 possible
mixture pairs
96.67% overlap
66.67% overlap
33.33 overlap
No overlap
Mixture Mixture
A
B
2.27x1014 possible
mixtures
D
Mixtures of 20
Mixture of 20
Mixtures of 10
A Mixture of 10
1.20x1023 possible
mixtures
Shared component
Mixtures of 30
Mixture of 30
1.54x1029 possible
mixtures
Component only in
mixture A
Component only in
mixture B
Not used in mixture
2.27x1032 possible
mixture pairs
3.24x1049 possible
mixture pairs
7.93x1056 possible
mixture pairs
1.10x1054 possible
mixture pairs
Used in mixture
Not used in mixture
Fig. 1. Odor mixtures used to test the resolution of the human olfactory
system. (A) Illustration of sample mixtures with exactly 10, 20, or 30 components
(green squares) picked from a collection of 128 odorous molecules (gray squares)
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and the number of possible mixtures of each type. (B) Example of one mixture
pair. (C to E) Schematics of each of the 13 types of odor pairs used for discrimination tests, along with the total number of possible mixture pairs of each type.
SCIENCE
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REPORTS
ture (9). The 128 molecules cover much of the
perceptual and physicochemical diversity of odorous molecules (1012) because the collection contains most of a collection of 86 odorous molecules
that were selected to be well distributed in both perceptual and physicochemical stimulus space (9).
To generate each mixture, we combined these
components together at equal ratios. The 128
components can be combined into 2.27 1014
different mixtures of exactly 10, 1.20 1023 different mixtures of exactly 20, and 1.54 1029
different mixtures of exactly 30 (Fig. 1A). The
most salient difference between two mixtures
with the same number of components is the percentage of components in which they differ.
We therefore performed psychophysical testing
to determine the resolution of the human olfactory system along this axis. We asked by what
percentage two mixtures must differ on average
so that they can be discriminated by the average
human nose. This percentage difference in components is the resolution of the olfactory system.
Subjects performed forced-choice discrimination tests to determine the discriminability of pairs
of mixtures (referred to here as mixture A and
mixture B) that varied in the percentage of shared
components (Fig. 1B). In double-blind experiments,
subjects were presented with three odor vials, two
Component in A and B
Component only in A
Component only in B
10 components
20 components
30 components
Mixture
A
Mixture
B
% mixture
90
overlap
60
30
95
75
50
25
100
% correct
80
60
40
Chance
20
Median
Subjects who can discriminate
Subjects who cannot discriminate
100
% correct
80
60
40
Chance
20
Median
Discriminable mixture
Non-discriminable mixture
Fig. 2. An empirical investigation of the resolution of the human olfactory system. (A) Schematic of the
discrimination tests carried out for mixtures of 10, 20, or 30 odorous molecules. (B and C) Results of discrimination
tests with 26 subjects asked to discriminate mixtures of 10 (left), 20 (middle), or 30 (right) components with decreasing
overlap from left to right. The dotted line represents the chance detection level (33.3%). For (B), dots represent
performance of individual subjects across 20 mixture pairs. For (C), dots represent average performance of all 26
subjects for a given mixture pair. Statistically significant discriminability (red dots) was assessed with a c2 test; P < 0.05.
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REPORTS
% mixture pairs
discriminable
50
Mixtures of 10
Mixtures of 20
Mixtures of 30
0
100
80
60
40
20
100
50
Mixtures of 10
Mixtures of 20
Mixtures of 30
0
100
# discriminable mixtures
100
80
% mixture overlap
60
40
20
1030
Mixtures of 10
Mixtures of 20
Mixtures of 30
1020
1010
100
0
20
40
60
80
100
% mixture overlap
allowing discrimination
% mixture overlap
s
# discriminable
mixtures
1030
1020
ate
Mixtures of 10
Mixtures of 20
Mixtures of 30
tim
s
se
ur
ixt
of
10
es
ur
ixt
20
of
es
ur
ixt
of
30
Smells
1010
Colors
Tones
100
0
20
40
60
80
100
104
% mixture overlap
allowing discrimination
108
106
1010
1012
1014
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Supplementary Materials
www.sciencemag.org/content/343/6177/1370/suppl/DC1
Materials and Methods
Fig. S1
Tables S1 and S2
2 December 2013; accepted 19 February 2014
10.1126/science.1249168
www.sciencemag.org