Springer

Handbook o
Odor
Buettner
Editor

123
Editor
Andrea Buettner
Fraunhofer Institute for Process Engineering and Packaging
Giggenhauser Str. 35
85354, Freising, Germany
andrea.buettner@ivv.fraunhofer.de

ISBN: 978-3-319-26930-6 e-ISBN: 978-3-319-26932-0

DOI 10.1007/978-3-319-26932-0
Springer Dordrecht Heidelberg London New York
Library of Congress Control Number: 2016953550

Springer International Publishing AG 2017

This work is subject to copyright. All rights are reserved by the Publisher,
whether the whole or part of the material is concerned, specifically
the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilm or in any other physical way, and
transmission or information storage and retrieval, electronic adaptation,
computer software, or by similar or dissimilar methodology now known or
hereafter developed.
The use of general descriptive names, registered names, trademarks,
service marks, etc. in this publication does not imply, even in the absence
of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice
and information in this book are believed to be true and accurate at the date
of publication. Neither the publisher nor the authors or the editors give a
warranty, express or implied, with respect to the material contained herein
or for any errors or omissions that may have been made.

Production and typesetting: le-tex publishing services GmbH, Leipzig

Typography and layout: schreiberVIS, Seeheim
Illustrations: Hippmann GbR, Schwarzenbruck
Cover design: eStudio Calamar Steinen, Barcelona
Cover production: WMXDesign GmbH, Heidelberg
Printing and binding: Printer Trento s.r.l., Trento

Printed on acid free paper

This Springer imprint is published by Springer Nature

The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham,
Switzerland

Matthias Laska
Human and A
Olfactory Capabilities Compared
Humans are traditionally considered to have
a poorly developed sense of smell that is clearly
inferior to that of nonhuman animals. This view,
however, is mainly based on an interpretation
of neuroanatomical and recent genetic findings,
and not on physiological or behavioral evidence.
An increasing number of studies now suggest that
the human sense of smell is much better than
previously thought and that olfaction plays a sig-
nificant role in regulating a wide variety of human
behaviors. This chapter, therefore, aims at sum-
marizing the current knowledge about human
olfactory capabilities and compares them to those
of animals.
Comparing olfactory capabilities between species is not
a trivial task. Several potentially confounding factors
have to be taken into consideration when trying to make
statements as to differences or similarities concerning
the efficiency of the sense of smell between species.
First, we find a high variability in results between
human studies. With regard to olfactory sensitivity, for
example, published mean threshold values for a given
odorant may vary up to 6 orders of magnitude [32.1].
This is probably, at least in part, due to different meth-
ods used to determine such thresholds. However, there
is a clear trend that more sophisticated psychophysi-
cal procedures employing signal detection methods and
rigorous stimulus control yield lower threshold values
than more simple procedures [32.2].
Second, the vast majority of psychophysical studies
on the human sense of smell only report mean values
of performance (usually plus a measure of variation),
but not the distribution of values or their range. This
is all the more problematic as measures of olfactory
performance are notorious for a high interindividual
variability. This is true for both studies on olfactory
sensitivity in which individual threshold values with
a given odorant have been reported to commonly vary
up to 3 orders of magnitude within a study popula-
tion [32.3], and for studies on olfactory discrimination
performance in which subjects have been found to vary
675
32. Human and Animal
32.1 Olfactory Sensitivity ............................. 678
32.2 Olfactory Discrimination Ability ............ 681
32.3 Qualitative Comparisons of Olfactory
Capabilities Between Species................ 684
32.3.1 Gathering Information
About the Chemical Environment 684
32.3.2 Foraging and Food Selection ...... 684
32.3.3 Spatial Orientation .................... 684
32.3.4 Social Communication ............... 684
32.3.5 Reproduction ............................ 685
32.3.6 Learning and Memory ................ 685
32.4 General Conclusions............................. 685
References................................................... 686
in their ability to distinguish between a given pair of
odorants from chance to perfect discrimination [32.4].
However, here too, more recent studies using state-of-
the-art psychophysical methods (pioneered by Cain and
coworkers) generally report a considerably lower range
between the most and least sensitive subjects within
a study population [32.5].
Third, it is inevitable to use different methods for
Part D | 32
assessing olfactory performance with animals and hu-
mans, and also with different species of animals. This
may affect the comparability of results. However, dif-
ferences between methods employed with different
species of animals are usually necessary adaptations for
meeting the physiological, anatomical, and behavioral
needs and limitations of the study species to success-
fully cooperate in a behavioral test. (In other words,
even if it was possible to use the same method with dif-
ferent species, this would very likely put one species
at an advantage over another species, thus invalidating
any comparisons. Therefore, it is better to aim for op-
timizing a testing method for each study species.) It is
commonly agreed that operant conditioning procedures
are the gold standard among the methods employed
with animals for assessing sensory performance [32.6],
and therefore only studies based on such procedures
are considered in this chapter. Figures 32.132.5 il-
lustrate operant conditioning procedures for assessing
 676 Part D Odorant Sensing and Physiological Effects

a) c) Fig. 32.1ad Olfactory conditioning

method used with spider monkeys.
(a) Portrait of a spider monkey
(Ateles geoffroyi). (b) The two-
choice apparatus used, viewed from
the animals side. It consists of
two manipulation boxes of which
one is baited with a Kelloggs
honeyloop while the other is empty,
depending on the odorant applied on
the absorbent paper strip attached
d) to the box. (d) A spider monkey
smelling at one of the absorbent
paper strips bearing an odorant used
either as a rewarded stimulus or as
b) an unrewarded stimulus. (c) A spider
monkey indicating its decision
for one of the two simultaneously
presented odorants by opening the
corresponding manipulation box
(courtesy of M. Laska)

a) c) Fig. 32.2ad Olfactory conditioning

method used with squirrel monkeys.
(a) Portrait of a squirrel monkey
(Saimiri sciureus). (b) Eppendorf
cups equipped with absorbent paper
strips. The Eppendorf cups serve as
Part D | 32

manipulation objects (artificial nuts)

that are either baited with a piece
of peanut or not, depending on the
odorant applied on the absorbent
paper strip. (d) Experimental setup.
d) An artificial nut tree is used to present
numerous artificial nuts, half of them
baited with a piece of peanut and
b) bearing an odorant used as rewarded
stimulus, and half of them empty
and bearing an odorant used as
unrewarded stimulus. (c) A squirrel
monkey inspecting an artificial nut
on a branch of the artificial nut tree
(courtesy of M. Laska)
 Human and Animal Olfactory Capabilities Compared Human and Animal Olfactory Capabilities Compared 677

a) b) Fig. 32.3ad Olfactory conditioning

method used with Asian elephants
(Elephas maximus). (a) Elephant
sniffing at the left odor port. (b) Ele-
phant sniffing at the right odor port.
(d) Elephant indicating its decision
for one of the two simultaneously
presented odorants by placing her
trunk onto the grid on top of the
corresponding odor port. (c) Elephant
receiving a carrot as a food reward
after a correct choice (courtesy of
M. Laska)

c) d)

a) b) Fig. 32.4ad Olfactory conditioning

method used with South African fur
seals. (a) Schematic drawing of the

Part D | 32
OP2
experimental set up. C: container
OP1 bearing an odor stimulus; V: ventilator
V
for ingoing airflow; O: outlet for
C outgoing airflow; SB: stimulus box;
OP1: odor port 1; OP2: odor port
SB 2. (b) Portrait of a South African
fur seal (Arctocephalus pusillus).
(c) Simultaneous presentation of two
c) d) odor stimuli to a fur seal. (d) A fur
seal sniffing at one of the two odor
ports (courtesy of M. Laska)
678 Part D Odorant Sensing and Physiological Effects
a) b)
Initiation Decision
S+ odor
Reward
Light barrier
Odor presentation
Odor S odor
applied
c) d)
olfactory performance in different species of mam-
mals.
Finally, animal studies usually only employ a low
number of individuals, sometimes only one or two an-
imals per species, thus making the use of mean values
arguable, and statements as to how representative the
findings are for the whole species difficult.
Despite all these difficulties, there are several good
reasons that make it worthwhile to compare olfactory
capabilities between humans and animals: first, such
comparisons allow us to study the neural and/or genetic
Part D | 32.1
32.1 Olfactory Sensitivity
The sensitivity of the sense of smell is usually as-
sessed by determining olfactory detection thresholds,
that is, the lowest concentration of a given odorant
that a human subject or an animal is able to de-
tect. Human olfactory detection thresholds have been
reported for a total of  3300 odorants [32.1]. In con-
trast, the total number of odorants tested in animals
is much lower. Table 32.1 summarizes the species
of mammals and the number of odorants for which
olfactory detection thresholds using operant condition-
ing procedures have been published. The total number
of species is 17 (please note that there exist about
 5500 species of mammals), and the highest num-
ber of odorants tested with a given species is 81.
These 17 species represent 7 of the 29 orders of mam-
mals.
Fig. 32.5ad Olfactory condition-
ing procedure used with mice.
(a) Schematic drawing of the exper-
imental setup. (b) A mouse (Mus
musculus) in front of the odor port.
(d) A mouse poking its head into the
odor port. (c) A mouse licking at the
water spout after a correct choice
(courtesy of M. Laska)
mechanisms underlying possible differences or simi-
larities in olfactory efficiency between species [32.7].
Second, between-species comparisons of olfactory per-
formance allow us to test hypotheses about the evo-
lution of sensory systems and the selective pressures
acting on them [32.8]. Finally, the integration of ani-
mal and human studies on olfactory performance may
help us to better understand medically relevant phe-
nomena such as aging processes or neurodegenerative
diseases, which are often accompanied by a loss of ol-
factory function [32.9, 10].
With the exception of four odorants tested with rats
(one explosive substance, and three explosive taggants,
that is, substances added to explosives to identify their
sources) and one odorant tested with mice (pyridazine),
all 138 odorants for which olfactory detection thresh-
olds have been determined with nonhuman mammals
have also been tested with human subjects, allowing for
direct comparisons of their performance.
In the following figures (as well as in all other
comparisons that follow), I compare the lowest mean
threshold values reported in human subjects to the
lowest individual threshold values reported in a given
animal species. The rationale for this is as follows:
1. Human studies only rarely report the range of
threshold values, but usually a mean value.
 Human and Animal Olfactory Capabilities Compared 32.1 Olfactory Sensitivity 679

Table 32.1 Animal species and number of odorants for which olfactory detection threshold values using operant condi-
tioning procedures have been published
No. Common name Scientific name Mammalian order Number of odorants tested
1 Spider monkey Ateles geoffroyi Primates 81
2 Mouse Mus musculus Rodentia 72
3 Squirrel monkey Saimiri sciureus Primates 61
4 Pigtail macaque Macaca nemestrina Primates 60
5 Rat Rattus norvegicus Rodentia 45
6 Short-tailed fruit bat Carollia perspicillata Chiroptera 18
7 Dog Canis lupus familiaris Carnivora 15
8 Common vampire bat Desmodus rotundus Chiroptera 15
9 Common mouse-eared bat Myotis myotis Chiroptera 13
10 Sea otter Enhydra lutris Carnivora 7
11 Pig Sus scrofa domestica Artiodactyla 5
12 Hedgehog Erinaceus europaeus Insectivora 4
13 Great fruit-eating bat Artibeus lituratus Chiroptera 3
14 Pale spear-nosed bat Phyllostomus discolor Chiroptera 3
15 Common shrew Sorex araneus Insectivora 3
16 Rabbit Oryctolagus cuniculus Lagomorpha 1
17 Harbor seal Phoca vitulina Carnivora 1

Fig. 32.6 Comparison of the olfactory

0 15
6
4 3 detection threshold values (expressed as vapor
4
1 10 8 Low sensitivity phase concentrations) of human subjects
6 13 3
15 1 2
6 5
3 10 for aliphatic n-carboxylic acids and those
2 10 6 1
8
4 10
8 15 14 2 2
of other mammalian species. Data points of
12 the human subjects (brown circles) represent
3 2
Threshold (log ppm)

3 1
2
1
4
12 3 the lowest mean threshold values reported in
4 12 7
the literature, and data points of all animal
2 12
4 1 species (numbered circles) represent the
5
2 lowest threshold values of individual animals
6 reported in the literature. Numbers in circles
High sensitivity

7
refer to the numbers assigned to each species
in Table 32.1 (human data: [32.1]; spider
7
8

Part D | 32.1
7 monkey data: [32.11]; mouse data: [32.12];
7 7 squirrel monkey data: [32.13]; pigtail macaque
9 7

7
data: [32.11]; rat data: [32.14]; short-tailed
10 fruit bat data: [32.15]; dog data: [32.16, 17];
Ethanoic n-Butanoic n-Hexanoic n-Octanoic common vampire bat data: [32.18]; sea otter
acid acid acid acid
data: [32.19]; hedgehog data: [32.20]; great
n-Propanoic n-Pentanoic n-Heptanoic
acid acid acid fruit-eating bat data: [32.18]; pale spear-nosed
bat data: [32.18]; common shrew data: [32.21])

2. Animal studies usually only employ a low num-
ber of individuals (in some cases only one animal),
making the use of mean values arguable.
3. Comparing the threshold value of the very best in-
dividual animal with the mean threshold value of
a group of human subjects minimizes the risk of un-
intentionally favoring humans over animals.
Figure 32.6 compares human olfactory detection
threshold values for aliphatic n-carboxylic acids to
those from other mammalian species. With the notable
exception of the dog, human subjects have a higher
sensitivity, that is, lower olfactory detection thresh-
olds, than the majority of mammalian species tested
with this class of odorants. (Please note that all data
points for the dog, except the one for n-heptanoic
acid, are from a study by Neuhaus [32.16] who em-
ployed only one animal. Later studies by other re-
searchers [32.17, 57] employed several dogs as well
as more rigorous stimulus control, and, interestingly,
680 Part D Odorant Sensing and Physiological Effects

Fig. 32.7 Comparison of the olfactory

3 9
5
9
detection threshold values (expressed as vapor
9

Low sensitivity
2 3 4 phase concentrations) of human subjects
for aliphatic alcohols and those of other
1
8 4 5 mammalian species. Data points of the
3
0 8 5
3 4 2 human subjects (brown circles) represent the
1
Threshold (log ppm)

8 1 lowest mean threshold values reported in

4 3
1 8 the literature, and data points of all animal
5 5 3
3
2 2
6
5 species (numbered circles) represent the
1 4 1 4
11 2 3 4 5 lowest threshold values of individual animals
3 reported in the literature. Numbers in circles
1 1

High sensitivity
4
refer to the numbers assigned to each species
in Table 32.1 (human data: [32.1]; spider
5 monkey data: [32.22]; mouse data: [32.2325];
11 squirrel monkey data: [32.26]; pigtail macaque
6
data: [32.26]; rat data: [32.27, 28]; short-tailed
2 fruit bat data: [32.15]; common vampire
7
Ethanol 1-Butanol 1-Hexanol 1-Octanol bat data: [32.18]; common mouse-eared bat
1-Propanol 1-Pentanol 1-Heptanol data: [32.29]; pig data: [32.30, 31])

Fig. 32.8 Comparison of the olfactory

2
4
Low sensitivity detection threshold values (expressed as vapor
1 3 phase concentrations) of human subjects for
aliphatic acetic esters and those of other
0
5 4 mammalian species. Data points of the
1
16
1 8 3
5
human subjects (brown circles) represent the
Threshold (log ppm)

1 3 1 10 4 6 lowest mean threshold values reported in

2 6
4 5
the literature, and data points of all animal
6 4 6 2
3 2 11 6 species (numbered circles) represent the
4
4 lowest threshold values of individual animals
4 1 3
1 1 3
reported in the literature. Numbers in circles
1 3
High sensitivity

5 3 refer to the numbers assigned to each species

2 5 in Table 32.1 (human data: [32.1]; spider
6
Part D | 32.1

7 monkey data: [32.32]; mouse data: [32.3335];

squirrel monkey data: [32.36]; pigtail macaque
7
data: [32.36]; rat data: [32.3739]; short-
8 tailed fruit bat data: [32.15]; dog data: [32.40];
Ethyl n-Butyl n-Hexyl iso-Pentyl common vampire bat data: [32.18]; sea
acetate acetate acetate acetate
otter data: [32.19]; pig data: [32.31]; rabbit
n-Propyl n-Pentyl iso-Butyl
acetate acetate acetate data: [32.41])

obtained markedly higher threshold values with these
odorants compared to those reported by Neuhaus.) The
mouse, another mammal with a reputation for a keen
sense of smell, is more sensitive than humans with
only three of the seven n-carboxylic acids whereas hu-
mans outperform this rodent with four of these seven
odorants.
Figure 32.7 compares human olfactory detection
threshold values for aliphatic 1-alcohols to those from
other mammalian species. Here, too, human subjects
have a higher sensitivity, that is, lower olfactory detec-
tion thresholds, than most of the other species tested.
It is interesting to note that humans outperform the rat,
another mammal believed to have a highly developed
sense of smell, with all seven 1-alcohols. Similarly, hu-
mans generally have lower olfactory detection thresh-
olds than the bats and nonhuman primates tested with
these odorants. The pig, in contrast, is clearly more sen-
sitive than humans with the two alcohols tested in this
species.
Figure 32.8 compares human olfactory detection
threshold values for aliphatic acetic esters to those from
 Human and Animal Olfactory Capabilities Compared 32.2 Olfactory Discrimination Ability 681

Animals Humans
Fig. 32.9 Comparison of all olfactory detection
more sensitive more sensitive threshold values between human subjects and
Spider monkey 18 57 animal species. Depicted are the number of odor-
Mouse 30 35 ants for which either human subjects or a given
Squirrel monkey 11 50 species of mammal are more sensitive (human
Pigtail macaque 6 54 data: [32.1]; spider monkey data: [32.11, 22, 25,
Rat 10 31
32, 4249]; mouse data: [32.12, 23, 25, 3335,
Short-tailed fruit bat 1 17
4244, 50, 51]; squirrel monkey data: [32.13, 26,
Dog 10 5
Common vampire bat 1 14
36, 4548]; pigtail macaque data: [32.11, 26, 36,
Common mouse-eared bat 0 13 4548]; rat data: [32.14, 27, 28, 3739, 45]; short-
Sea otter 0 7 tailed fruit bat data: [32.15]; dog data: [32.16, 17,
Pig 2 3 40, 5254]; common vampire bat data: [32.18];
Hedgehog 1 3 common mouse-eared bat data: [32.29]; sea otter
Great fruit-eating bat 1 2 data: [32.19]; pig data: [32.30, 31, 55]; hedgehog
Pale spear-nosed bat 1 2 data: [32.20]; great fruit-eating bat data: [32.18];
Common shrew 0 3
pale spear-nosed bat data: [32.18]; common shrew
Rabbit 0 1
data: [32.21]; rabbit data: [32.41]; harbor seal
Harbor seal 1 0
data: [32.56])
40 30 20 10 0 10 20 30 40 50 60
Number of odorants

other mammalian species. With only two exceptions
(spider monkeys and squirrel monkeys with n-butyl ac-
etate), human subjects have lower olfactory detection
thresholds, that is, a higher sensitivity for these odorants
than all other mammal species tested, including dogs,
mice, and rats. With only few exceptions, humans are
also more sensitive for aliphatic acetic esters than spi-
der monkeys, squirrel monkeys, and pigtail macaques.
This is remarkable given that these nonhuman primates
are highly frugivorous suggesting that a high olfactory
sensitivity for fruit-associated odorants such as acetic
esters should be adaptive for these species. However,
both human and nonhuman primates generally outper-
form granivorous species, such as rats, insectivorous
lower olfactory detection thresholds, that is, a higher
sensitivity with the majority of odorants tested so far
compared to all other mammal species tested so far.
This includes species traditionally considered to have
a highly developed sense of smell, such as mice, rats,
hedgehogs, shrews, pigs, and rabbits.
It is interesting to note that humans outperform
even the dog, often considered as the undisputed super-
nose of the animal kingdom, with 5 of the 15 odorants
tested with both species. The fact that these 5 odorants
comprise plant odor components such as -ionone and
n-pentyl acetate suggests that the behavioral relevance
of odorants rather than neuroanatomical or genetic fea-
tures may strongly affect a species olfactory sensitivity.

species such as bats, and herbivorous species such as
the rabbit with this class of odorants.
Figure 32.9 summarizes all comparisons of olfac-
tory detection thresholds between human subjects and
other mammal species. Depicted are the number of
odorants for which either human subjects or a given
species of mammal are more sensitive. With the excep-
tion of the dog (and the harbor seal, which has been
tested with only one odorant), human subjects have
Part D | 32.2
This idea is further supported by the fact that 7 of the
10 odorants for which the dog has been reported to be
more sensitive than humans comprise carboxylic acids
and thus typical components of the odor of prey species
of the dog.
Thus, based on these comparisons, and contrary to
traditional textbook wisdom, humans are not generally
inferior in their olfactory sensitivity compared to ani-
mals.

32.2 Olfactory Discrimination Ability

Olfactory discrimination can be defined as the abil-
ity to reliably respond differently to the successive
presentation of two nonidentical odorants. Thus, ol-
factory discrimination is usually assessed by deter-
mining the proportion of correct responses with re-
peated presentations of a given pair of odorants.
A statistical criterion can then be used to decide
whether a human subject or an animal is able to dis-
criminate between the two odorants in question or
not.
Human studies on olfactory discrimination ca-
pabilities usually either employ structurally related
682 Part D Odorant Sensing and Physiological Effects

Table 32.2 Between-species comparison of olfactory discrimination performance with aliphatic odorants sharing the
same functional group but differing in carbon chain length
Human Squirrel Asian Fur seals CD-1 mice Honey bees
subjects monkeys elephants
1-Alcohols +++++ + +++ +++++ ++++++ +++++
n-Aldehydes ++++ +++ +++ +++++ ++++++ ++++++
2-Ketones +++++ +++ +++ +++++ ++++++ ++++++
Acetic esters +++++ +++ +++ ++++ ++++++
n-Carboxylic acids ++++++ +++ +++ +++++ ++++++
Success rate 25/30 13/15 15/15 24/25 30/30 17/18
A + symbol indicates that the group of human subjects or animals succeeded in discriminating a given aliphatic odor pair, a 
symbol indicates failure to do so, and a symbol indicates that this odor pair was not tested. The six symbols in each table cell
refer to the discrimination of carbon chain lengths C4 versus C5 , C4 versus C6 , C4 versus C7 , C5 versus C6 , C5 versus C7 , and C6
versus C7 , respectively. Human data: [32.5860]; squirrel monkey data: [32.59, 61, 62]; Asian elephant data: [32.63, 64]; fur seal
data: [32.65]; mouse data: [32.66]; honeybee data: [32.67]

Table 32.3 Between-species comparison of olfactory discrimination performance with aliphatic odorants sharing the
same carbon length but differing in functional group
Human subjects Squirrel monkeys CD-1 mice Honey bees
1-Alcohols versus n-aldehydes +++ +++ +++ +++
1-Alcohols versus 2-ketones +++ +++ +++ +++
1-Alcohols versus n-carboxylic acids +++ +++ +++
n-Aldehydes versus 2-ketones +++ +++ +++ +++
n-Aldehydes versus n-carboxylic acids +++ +++ +++
2-Ketones versus n-carboxylic acids +++ +++ +++
Success rate 18/18 18/18 18/18 9/9
A + symbol indicates that the group of human subjects or animals succeeded in discriminating a given aliphatic odor pair, a 
symbol indicates failure to do so, and a symbol indicates that this odor pair was not tested. The three symbols in each table cell
refer to the discrimination of odorants that share chain lengths of either 4, or 6, or 8 carbon atoms, respectively. Human data: [32.68];
squirrel monkey data: [32.69]; mouse data: [32.66]; honeybee data: [32.67]

monomolecular odorants (to investigate correlations be- ants, the proportion of successfully discriminated odor
tween structure and perceived quality of odorants), or pairs is slightly lower in humans compared to squirrel
complex odor mixtures of commercial use such as fra- monkeys, fur seals, honey bees, Asian elephants, and
Part D | 32.2

grances (perfumes, body care products) or food odors
(wines, coffees, artificial flavors). In contrast, the vast
majority of animal studies assessing olfactory discrimi-
nation capabilities employ naturally occurring complex
odor mixtures that are behaviorally relevant for the
species under study such as conspecific body odors,
species-typical food odors, or predator odors. As a con-
sequence, there is only little overlap in the stimuli used
between human and animal studies on olfactory dis-
crimination. However, at least a few animal species
have also been studied for their ability to distinguish
between some of the structurally related monomolecu-
lar odorants tested with humans.
Table 32.2 compares the ability of humans and sev-
eral species of animals to discriminate between mem-
bers of homologous series of aliphatic odorants. These
are odorants sharing the same oxygen-containing func-
tional group (e.g., an alcohol- or an aldehyde group) but
differing in carbon chain length. With this set of odor-
mice. Nevertheless, humans succeeded with more than
80% of these odor pairs, which are both structurally and
qualitatively similar to each other.
Table 32.3 compares the ability of humans and
several species of animals to discriminate between
aliphatic odorants sharing the same carbon chain length
but differing in functional group. With this set of odor-
ants, not only squirrel monkeys, mice, and honey bees,
but also human subjects successfully discriminated be-
tween all odor pairs tested.
Table 32.4 compares the ability of humans and
several species of animals to discriminate between
enantiomers. These are pairs of molecules with mirror-
image structures that exhibit identical chemical and
physical properties except for their optical activity, that
is, rotation of polarized light. They are particularly
useful for assessing how molecular structure is en-
coded by olfactory systems as perceptual differences
between enantiomers cannot be caused by differing
 Human and Animal Olfactory Capabilities Compared 32.2 Olfactory Discrimination Ability 683

Table 32.4 Between-species comparison of olfactory discrimination performance with enantiomers

Human Squirrel Asian Fur CD-1 Honey Pigtail SD/LE
subjects monkeys elephants seals mice bees macaques rats
Carvone + + + + + + + +
Dihydrocarvone + + + + + +
Dihydrocarveol + + + + + +
Limonene + + +  + + + +
-Pinene + + + + +
Isopulegol   +  + +
Menthol   + + + +
-Citronellol   + + + +
Rose oxide   +  + 
Fenchone  + + + +  +
Limonene oxide   + + + 
Camphor   +  + 
Success rate 5/12 6/12 12/12 8/12 11/11 5/8 5/6 3/3
A + symbol indicates that the group of animals or subjects succeeded in discriminating a given enantiomeric odor pair, and a 
symbol indicates failure to do so. Human data: [32.70, 71]; squirrel monkey data: [32.72, 73]; Asian elephant data: [32.63]; fur seal
data: [32.74]; mouse data: [32.75]; honeybee data: [32.76]; pigtail macaque data: [32.73]; rat data: [32.77, 78]

diffusion rates in the mucus covering the olfactory ep-
ithelium or differing airmucus partition coefficients,
but must originate from chiral selectivity at the receptor
level [32.79].
Human subjects, as a group, are only able to dis-
criminate between 5 of the 12 enantiomeric odor pairs
tested and thus perform similar to squirrel monkeys,
which succeeded with 6 out of 12 pairs. Asian ele-
phants, mice, and rats, in contrast, succeeded with 12
out of 12, 11 out of 11, and 3 out of 3 of these odor pairs.
South African fur seals, pigtail macaques, and honey-
bees are able to discriminate between the majority, but
not all enantiomeric odor pairs tested with these species
(Table 32.4).
to be reliably perceived as stronger than the stan-
dard.
The human JND has been found to be odorant-
dependent and Weber fractions have been reported
to range from 0:09 for n-pentyl butyrate, 0:16 for
n-pentanol, 0:30 for pyridine, 0:35 for n-butanol, to
0:47 for phenylethanol [32.8082]. The only study that
directly compared JNDs between species found the We-
ber fraction for n-pentyl acetate to be 0:041 in rats and
thus considerably lower than the 0:315 found with this
odorant in humans [32.83]. However, a later study re-
ported the rats Weber fraction for the same odorant to
be 0:28, and thus comparable to that of humans [32.84].
A possible explanation for the extreme paucity of

Based on these comparisons, humans appear to
have a slightly inferior olfactory discrimination ability
compared to some species such as mice and Asian ele-
phants. However, their performance in discriminating
between structurally related monomolecular odorants
appears to be comparable to that of squirrel monkeys,
fur seals, and honeybees.
A special, and only rarely investigated, aspect of
olfactory discrimination performance is the ability to
distinguish between different concentrations of a given
odorant. The smallest concentration difference that
a nose can reliably detect is often referred to as just
noticeable difference (JND) and is usually expressed
as a so-called Weber fraction (according to Webers
law, which states that the JND between two stimuli is
proportional to the magnitude of the stimuli:  I=I D
constant). A Weber fraction of 0:3, for example, in-
dicates that a stimulus has to be presented at a 30%
higher intensity, relative to a standard stimulus, in order
Part D | 32.2
animal data on olfactory JNDs is their difficulty in
learning the concept that two different concentrations
of the same odorant should have different reward val-
ues. To understand this difficulty, one must know that
in operant conditioning procedures an animal learns
that an odorant A is rewarded and that an odorant B
(or a blank stimulus) is not rewarded. Once an ani-
mal has successfully learned the association between
odorant A and a reward, it will consider this odorant
as a rewarded stimulus irrespective of its concentration.
Biologically, this makes perfect sense as odors that are
behaviorally relevant for an animal hardly ever change
their significance as a function of concentration: a food
odor will always be attractive, whether detected at high
or low concentrations, and a predator odor will be al-
ways avoided, whether at high or low concentrations.
Thus, it takes an animal extensive training to overcome
this perseverance with regard to the learned reward
value of odorants.
684 Part D Odorant Sensing and Physiological Effects
32.3 Qualitative Comparisons of Olfactory Capabilities
Between Species
In addition to using quantifiable measures of olfac-
tory performance, such as sensitivity and discrimination
ability, one can also try to compare the sense of smell
between species using qualitative measures. In the sim-
plest case, this means to assess whether a given species
possesses a certain ability or not. To this end, it might be
useful to take a look at different behavioral contexts in
which the sense of smell is known to play a role for an-
imals and then to ask whether human subjects are able
to use their noses for the same purpose.
32.3.1 Gathering Information
About the Chemical Environment
The ability to gather information about the chemical en-
vironment is almost ubiquitous in the animal kingdom.
In most species of animals, the detection of chemi-
cal hazards, for example, leads to adaptive behavioral
responses intended to minimize further exposure. Hu-
mans are no exception to this rule and display adaptive
behaviors such as head turning, eye closure, apnea (sus-
pension of breathing), sneezing, and coughing when
exposed to harmful volatile chemicals. Although some
of these reflex-like responses involve the nasal trigem-
inal system, the significance of the olfactory system in
this context becomes apparent when considering anos-
mic subjects: humans without a functioning sense of
smell run a significantly higher risk of suffering from
gas poisoning [32.85] and of not detecting fire [32.86]
compared to healthy controls.
Part D | 32.3
32.3.2 Foraging and Food Selection
The ability to find and select food using the sense
of smell is also widespread among animal species.
Human babies, similar to the offspring of other mam-
mals, already display adaptive behavioral responses
such as positive head turning and gaping mouth move-
ments when presented with the odor of their mothers
breast or the odor of breast milk [32.87]. Although hu-
mans nowadays hardly ever need to use their noses
to forage, that is, to search for food, they do possess
the ability to follow a food odor scent trail [32.88].
The involvement of the sense of smell in human food
selection is more obvious: anosmic subjects run a sig-
nificantly higher risk of suffering from food poisoning
compared to healthy controls with an intact sense of
smell [32.89]. Similarly, the risk of suffering from
malnutrition is markedly higher in anosmic subjects
compared to normosmic control subjects [32.90]. Not
surprisingly, anosmic subjects particularly those who
lost their sense of smell instantaneously, for exam-
ple, due to head trauma often report a consider-
able loss of quality of life with regard to enjoying
food [32.91].
32.3.3 Spatial Orientation
Quite a number of animal species rely on their sense
of smell for spatial orientation. This is particularly true
for nocturnal and subterranean species. There are two
basic mechanisms that allow animals to find their way
through their habitat using their sense of smell: they
can either use existing landmarks that emit an odor,
or they can deposit scent marks themselves at certain
points in their habitat. Both types of odor sources serve
as navigational landmarks that can be used by animals
to recognize their position in space.
Although humans with an intact sense of vision
hardly ever need to use their noses for finding their way,
they do possess the ability to follow scent trails laid
by conspecifics and when blindfolded [32.88]. A re-
cent study suggests the existence of spatial information
processing in the human olfactory system and thus of
an implicit ability for directional smelling [32.92]. Fur-
ther, there is anecdotal evidence that blind persons use
olfactory landmarks as sensory cues for spatial orienta-
tion [32.93, 94].
32.3.4 Social Communication
Many species of animals have been shown to use body-
borne odors for social communication. Such odors may
convey information about species, social group, sex,
age, reproductive status, health status, social rank, in-
dividual identity, genetic relatedness, dietary habit, and
emotional state of the odor donor.
Psychophysical studies have demonstrated that hu-
mans are able to correctly assign body odors to
sex [32.95] and to age classes [32.96]. Similarly, it has
been shown that humans are able to correctly assign
body odors to reproductive status [32.97, 98] and to
health status [32.99, 100].
Further, humans are able to distinguish between in-
dividual body odors [32.101] and between body odors
of kin and nonkin [32.102, 103]. This includes mutual
recognition of the body odors between mothers and
their babies [32.104].
Humans are also able to distinguish between the
body odors of vegetarians and meat eaters [32.105] and
 Human and Animal Olfactory Capabilities Compared
thus they can sniff out the dietary habits of conspecifics.
Recent studies suggest that humans are also able to de-
tect emotions via body odors [32.106, 107].
Dogs, mice, and rats are not only able to discrimi-
nate between the odors of individual conspecifics, but
also between the odors of individuals of other species,
for example, of individual humans. However, it has been
demonstrated that human subjects are also able to cor-
rectly identify the odor of their own pet dog among other
dog odor samples [32.108], to correctly assign body
odor samples to individual gorillas [32.109], and to cor-
rectly discriminate between the body odors of mouse
strains that only differ from each other in their major his-
tocompatibility complex (MHC) genes [32.110].
32.3.5 Reproduction
The ability to correctly identify the reproductive status
of potential mates using olfactory cues is widespread
among mammals. Similarly, mate choice has been
shown to be based on or at least influenced by ol-
factory cues in a number of mammal species. Human
males are able to distinguish between female body
odors from different phases of the estrous cycle and pre-
fer the body odor of females produced around the time
of ovulation [32.97, 98, 111]. Several lines of evidence
suggest that human mate choice may be influenced
by body odors and that, as is the case in mice, MHC
genes are involved in the formation of individual odor
signatures [32.112114]. A recent study found that
congenitally anosmic men exhibit a strongly reduced
32.4 General Conclusions
The amount of published data on quantifiable measures
of olfactory performance, which allow for direct com-
parisons between humans and animals, is rather limited.
However, based on this limited set of data the following
conclusions can be drawn:
Human subjects have lower olfactory detection
thresholds, that is, a higher sensitivity with the major-
ity of odorants tested so far compared to most of the
mammal species tested so far. This includes species tra-
ditionally considered to have a highly developed sense
of smell such as mice, rats, hedgehogs, shrews, pigs,
and rabbits.
Human subjects appear to have a slightly inferior ol-
factory discrimination ability compared to species such
as mice and Asian elephants. However, their perfor-
32.4 General Conclusions 685
number of sexual relationships compared to norm-
osmic men, and that congenitally anosmic women feel
less secure about their romantic partner compared to
normosmic women [32.115]. Further, anosmia-related
depression has been shown to reduce sexual appetite in
humans [32.116].
32.3.6 Learning and Memory
Learning and memory are inevitably linked to sensory
input. Many species of animals rely on olfactory cues
for learning about their environment or about situa-
tional contexts as well as for building and retrieving
memories. Humans are no exception to this as they
are able to rapidly and robustly learn long-lasting as-
sociations between food odors and positive or negative
physiological consequences of ingestion [32.117]. Sim-
ilarly, humans are able to rapidly and robustly learn
appetitive and aversive associations between odors and
visual stimuli [32.118]. Several studies reported human
long-term memory for odors to be outstanding and su-
perior to that for other sensory modalities [32.119]. The
longest interval tested so far with humans for successful
odor recognition was 1 year [32.120], and for above-
chance level retention of odorname associations even
9 years [32.121].
From these qualitative comparisons of olfactory ca-
pabilities, one must conclude that humans have at least
the basic ability of using their sense of smell in all be-
havioral contexts in which animals are known to use
their noses.
Part D | 32.4
mance in discriminating between structurally related
monomolecular odorants appears to be comparable to
that of species such as squirrel monkeys, fur seals, and
honeybees.
Qualitative comparisons of olfactory capabilities
suggest that human subjects have at least the basic
ability of using their sense of smell in all behavioral
contexts in which animals are known to use their noses.
This includes gathering of information about the chem-
ical environment, food selection, spatial orientation,
social communication, reproduction, as well as learn-
ing and memory.
Taken together, these findings suggest that the human
sense of smell is not generally inferior compared to that
of animals and much better than traditionally thought.
686 Part D Odorant Sensing and Physiological Effects
References
32.1 L.J. van Gemert: Odour Thresholds. Compilations
of Odour Threshold Values in Air, Water and Other
Media, 2nd edn. (OPP, Utrecht 2011)
32.2 R. Schmidt, W.S. Cain: Making scents: Dynamic
olfactometry for threshold measurement, Chem.
Senses 35, 109120 (2010)
32.3 J.C. Stevens, W.S. Cain, R.J. Burke: Variability of
olfactory thresholds, Chem. Senses 13, 643653
(1988)
32.4 M. Laska, A. Ringh: How big is the gap between
olfactory detectin and recognition of aliphatic
aldehydes?, Attent. Percept. Psychophys. 72,
806812 (2010)
32.5 J.E. Cometto-Muiz, M.H. Abraham: Structure-
activity relationships on the odor detectability
of homologous carboxylic acids by humans, Exp.
Brain Res. 207, 7584 (2010)
32.6 J.M. Pearce: Animal Learning and Cognition (Psy-
chology, New York 2008)
32.7 B.W. Ache, J.M. Young: Olfaction: Diverse species,
conserved principles, Neuron 48, 417430 (2005)
32.8 R.A. Barton: Olfactory evolution and behavioral
ecology in primates, Am. J. Primatol. 68, 545558
(2006)
32.9 R.L. Doty, I. Petersen, N. Mensah, K. Christensen:
Genetic and environmental influences on odor
identification ability in the very old, Psychol. Ag-
ing 26, 864871 (2011)
32.10 M. Barresi, R. Ciurleo, S. Giacoppo, V.F. Cuzzola,
D. Celi, P. Bramanti, S. Marino: Evaluation of
olfactory dysfunction in neurodegenerative dis-
eases, J. Neurol. Sci. 323, 1624 (2012)
32.11 M. Laska, A. Wieser, R.M. Rivas Bautista, L.T. Her-
nandez Salazar: Olfactory sensitivity for carboxylic
acids in spider monkeys and pigtail macaques,
Chem. Senses 29, 101109 (2004)
32.12 S.C. Gven, M. Laska: Olfactory sensitivity and
Part D | 32
odor structure-activity relationships for aliphatic
carboxylic acids in CD-1 mice, PLoS ONE 7, e34301
(2012)
32.13 M. Laska, A. Seibt, A. Weber: Microsmatic primates
revisited Olfactory sensitivity in the squirrel
monkey, Chem. Senses 25, 4753 (2000)
32.14 B.M. Slotnick, G.A. Bell, H. Panhuber, D.G. Laing:
Detection and discrimination of propionic acid
after removal of its 2-DG identified major focus
in the olfactory bulb: A psychophysical analysis,
Brain Res. 762, 8996 (1997)
32.15 M. Laska: Olfactory sensitivity to food odor com-
ponents in the short-tailed fruit bat, Carollia per-
spicillata (Phyllostomatidae, Chiroptera), J. Comp.
Physiol. A 166, 395399 (1990)
32.16 W. Neuhaus: ber die Riechschrfe des Hundes
fr Fettsuren, Z. Vergl. Physiol. 35, 527552 (1953)
32.17 D.G. Moulton, D.H. Ashton, J.T. Eayrs: Studies in
olfactory acuity. 4. Relative detectability of n-
aliphatic acids by the dog, Anim. Behav. 8, 117
128 (1960)
32.18 U. Schmidt: Vergleichende Riechschwellenbes-
timmungen bei neotropischen Chiropteren
(Desmodus rotundus, Artibeus lituratus, Phyl-
lostomus discolor), Z. Sugetierkd. 40, 269298
(1975)
32.19 J. Hammock: Structure, Function and Context: The
Impact of Morphometry and Ecology on Olfactory
Sensitivity, Ph.D. Thesis (MIT, Cambridge 2005)
32.20 H. Bretting: Die Bestimmung der Riechschwellen
bei Igeln fr Einige Fettsuren, Z. Sugetierkd. 37,
286311 (1972)
32.21 L. Sigmund, F. Sedlacek: Morphometry of the ol-
factory organ and olfactory thresholds of some
fatty acids in Sorex araneus, Acta Zool. Fennica
173, 249251 (1985)
32.22 M. Laska, R.M. Rivas Bautista, L.T. Hernandez
Salazar: Olfactory sensitivity for aliphatic alcohols
and aldehydes in spider monkeys, Ateles geof-
froyi, Am. J. Phys. Anthropol. 129, 112120 (2006)
32.23 J. Larson, J.S. Hoffman, A. Guidotti, E. Costa: Ol-
factory discrimination learning in heterozygous
reeler mice, Brain Res. 971, 4046 (2003)
32.24 D.W. Smith, S. Thach, E.L. Marshall, M.G. Men-
doza, S.J. Kleene: Mice lacking NKCC1 have normal
olfactory sensitivity, Physiol. Behav. 93, 4449
(2008)
32.25 P.K. Ltvedt, S.K. Murali, L.T. Hernandez Salazar,
M. Laska: Olfactory sensitivity for green odors
(aliphatic C6 alcohols and C6 aldehydes) A com-
parative study in male CD-1 mice (Mus musculus)
and female spider monkeys (Ateles geoffroyi),
Pharmacol. Biochem. Behav. 101, 450457 (2012)
32.26 M. Laska, A. Seibt: Olfactory sensitivity for
aliphatic alcohols in squirrel monkeys and pig-
tail macaques, J. Exp. Biol. 205, 16331643 (2002)
32.27 D.G. Moulton, J.T. Eayrs: Studies in olfactory acu-
ity. 2. Relative detectability of n-aliphatic alco-
hols by the rat, Q. J. Exp. Psychol. 12, 99109
(1960)
32.28 D.G. Laing: A comparative study of the olfactory
sensitivity of humans and rats, Chem. Senses Fla-
vor 1, 257269 (1975)
32.29 U. Schmidt, C. Schmidt: Olfactory thresholds in
four microchiropteran bat species, Proc. 4th Int.
Bat Res. Conf., Nairobi (1978) pp. 713
32.30 J.B. Jones, C.M. Wathes, K.C. Persaud, R.P. White,
R.B. Jones: Acute and chronic exposure to ammo-
nia and olfactory acuity for n-butanol in the pig,
Appl. Anim. Behav. Sci. 71, 1328 (2001)
32.31 L.M. Sndergaard, I.E. Holm, M.S. Herskin, F. Dag-
naes-Hansen, M.G. Johansen, A.L. Jrgensen,
J. Ladewig: Determination of odor detection
threshold in the Gttingen minipig, Chem. Senses
35, 727734 (2010)
32.32 L.T. Hernandez Salazar, M. Laska, E. Rodriguez
Luna: Olfactory sensitivity for aliphatic esters in
spider monkeys, Ateles geoffroyi, Behav. Neu-
rosci. 117, 11421149 (2003)
32.33 V. Vedin, B. Slotnick, A. Berghard: Zonal ablation
of the olfactory sensory neuroepithelium of the
mouse: Effects on odorant detection, Eur. J. Neu-
rosci. 20, 18581864 (2004)

32.34 J.C. Walker, R.J. OConnell: Computerized odor
psychophysical testing in mice, Chem. Senses 11,
439453 (1986)
32.35 A.C. Clevenger, D. Restrepo: Evaluation of the va-
lidity of a maximum likelihood adaptive staircase
procedure for measurement of olfactory detection
threshold in mice, Chem. Senses 31, 926 (2006)
32.36 M. Laska, A. Seibt: Olfactory sensitivity for
aliphatic esters in squirrel monkeys and pigtail
macaques, Behav. Brain Res. 134, 165174 (2002)
32.37 S. Krmer, R. Apfelbach: Olfactory sensitivity,
learning and cognition in young adult and aged
male Wistar rats, Physiol. Behav. 81, 435442
(2004)
32.38 S. Pierson: Conditioned suppression to odorous
stimuli in the rat, J. Comp. Physiol. Psychol. 86,
708717 (1974)
32.39 R.G. Davis: Olfactory psychophysical parameters
in man, rat, dog and pigeon, J. Comp. Physiol.
Psychol. 85, 221232 (1973)
32.40 D.B. Walker, J.C. Walker, P.J. Cavnar, J.L. Tay-
lor, D.H. Pickel, S.B. Hall, J.C. Suarez: Naturalis-
tic quantification of canine olfactory sensitivity,
Appl. Anim. Behav. Sci. 97, 241254 (2006)
32.41 D.G. Moulton, A. Celebi, R.P. Fink: Olfaction in
mammals two aspects: Proliferation of cells
in the olfactory epithelium and sensitivity to
odours. In: Ciba Foundation Symposium on Taste
and Smell in Vertebrates, ed. by G.E.W. Wolsten-
holme, J. Knight (Churchill, London 1970)
32.42 D. Joshi, M. Vlkl, G.M. Shepherd, M. Laska: Olfac-
tory sensitivity for enantiomers and their racemic
mixtures A comparative study in CD-1 mice
and spider monkeys, Chem. Senses 31, 655664
(2006)
32.43 M. Laska, O. Persson, L.T. Hernandez Salazar: Ol-
factory sensitivity for alkylpyrazines A compar-
ative study in CD-1 mice and spider monkeys,
J. Exp. Zool. A 311, 278288 (2009)
32.44 H. Walln, I. Engstrm, L.T. Hernandez Salazar,
M. Laska: Olfactory sensitivity for six amino acids:
A comparative study in CD-1 mice and spider
monkeys, Amino Acids 42, 14751485 (2012)
32.45 M. Laska, M. Fendt, A. Wieser, T. Endres, L.T. Her-
nandez Salazar, R. Apfelbach: Detecting danger
or just another odorant? Olfactory sensitivity for
the fox odor component 2,4,5-trimethyl thiazo-
line in four species of mammals, Physiol. Behav.
84, 211215 (2005)
32.46 M. Laska, A. Wieser, L.T. Hernandez Salazar: Ol-
factory responsiveness to two odorous steroids
in three species of nonhuman primates, Chem.
Senses 30, 505511 (2005)
32.47 M. Laska, D. Hfelmann, D. Huber, M. Schu-
macher: Does the frequency of occurrence of
odorants in the chemical environment deter-
mine olfactory sensitivity? A study with acyclic
monoterpene alcohols in three species of nonhu-
man primates, J. Chem. Ecol. 32, 13171331 (2006)
32.48 M. Laska, R.M. Rivas Bautista, D. Hfelmann,
V. Sterlemann, L.T. Hernandez Salazar: Olfactory
sensitivity for putrefaction-associated thiols and
Human and Animal Olfactory Capabilities Compared References 687
indols in three species of nonhuman primates,
J. Exp. Biol. 210, 41694178 (2007)
32.49 L. Kjeldmand, L.T. Hernandez Salazar, M. Laska:
Olfactory sensitivity for sperm-attractant aro-
matic aldehydes A comparative study in human
subjects and spider monkeys, J. Comp. Physiol. A
197, 1523 (2011)
32.50 M. Laska, D. Joshi, G.M. Shepherd: Olfactory sen-
sitivity for aliphatic aldehydes in CD-1 mice, Be-
hav. Brain Res. 167, 349354 (2006)
32.51 L. Larsson, M. Laska: Ultra-high olfactory sensi-
tivity for the human sperm-attractant aromatic
aldehyde bourgeonal in CD-1 mice, Neurosci. Res.
71, 355360 (2011)
32.52 W. Neuhaus: Die Riechschwellen des Hundes fr
Jonon und thylmercaptan und ihr Verhltnis zu
anderen Riechschwellen bei Hund und Mensch,
Z. Naturforsch. 9, 560567 (1954)
32.53 P.I. Ezeh, L.J. Myers, L.A. Hanrahan, R.J. Kemp-
painen, K.A. Cummins: Effects of steroids on the
olfactory function of the dog, Physiol. Behav. 51,
11831187 (1992)
32.54 L.J. Myers, R. Pugh: Threshold of the dog for
detection of inhaled eugenol and benzaldehyde
determined by electroencephalographic and be-
havioral olfactometry, Am. J. Vet. Res. 46, 2409
2412 (1985)
32.55 K.M. Dorries, E. Adkins-Regan, B.P. Halpern:
Olfactory sensitivity to the pheromone an-
drostenone in sexually dimorphic in the pig,
Physiol. Behav. 57, 255259 (1995)
32.56 S. Kowalewsky, M. Dambach, B. Mauck, G. Dehn-
hardt: High olfactory sensitivity for dimethyl sul-
phide in harbour seal, Biol. Lett. 2, 106109
(2006)
32.57 D.A. Marshall, L. Blumer, D.G. Moulton: Odor de-
tection curves for n-pentanoic acid in dogs and
humans, Chem. Senses 6, 445453 (1981)
32.58 M. Laska, P. Teubner: Odor structure-activity rela-
tionships of carboxylic acids correspond between
Part D | 32
squirrel monkeys and humans, Am. J. Physiol.
274, R1639R1645 (1998)
32.59 M. Laska, P. Teubner: Olfactory discrimination
ability for homologous series of aliphatic alcohols
and aldehydes, Chem. Senses 24, 263270 (1999)
32.60 M. Laska, F. Hbener: Olfactory discrimination
ability for homologous series of aliphatic ketones
and acetic esters, Behav. Brain Res. 119, 193201
(2001)
32.61 M. Laska, S. Trolp, P. Teubner: Odor structure-ac-
tivity relationships compared in human and non-
human primates, Behav. Neurosci. 113, 981007
(1999)
32.62 M. Laska, D. Freyer: Olfactory discrimination abil-
ity for aliphatic esters in squirrel monkeys and
humans, Chem. Senses 22, 457465 (1997)
32.63 A. Rizvanovic, M. Amundin, M. Laska: Olfactory
discrimination ability of Asian elephants (Elephas
maximus) for structurally related odorants, Chem.
Senses 38, 107118 (2013)
32.64 J. Arvidsson, M. Amundin, M. Laska: Successful
acquisition of an olfactory discrimination test by
688 Part D Odorant Sensing and Physiological Effects
Asian elephants, Elephas maximus, Physiol. Be-
hav. 105, 809814 (2012)
32.65 M. Laska, E. Lord, S. Selin, M. Amundin: Olfac-
tory discrimination of aliphatic odorants in South
African fur seals (Arctocephalus pusillus), J. Comp.
Psychol. 124, 187193 (2010)
32.66 M. Laska, . Rosandher, S. Hommen: Olfactory
discrimination of aliphatic odorants at 1 ppm: Too
easy for CD-1 mice to show odor structure-activity
relationships?, J. Comp. Physiol. A 194, 971980
(2008)
32.67 M. Laska, C.G. Galizia, M. Giurfa, R. Menzel: Ol-
factory discrimination ability and odor structure-
activity relationships in honeybees, Chem. Senses
24, 429438 (1999)
32.68 M. Laska, S. Ayabe-Kanamura, F. Hbener,
S. Saito: Olfactory discrimination ability for
aliphatic odorants as a function of oxygen moiety,
Chem. Senses 25, 189197 (2000)
32.69 M. Pontz: Untersuchung des geruchlichen Un-
terscheidungsvermgens von Totenkopfaffen
(Saimiri sciureus) fr aliphatische Substanzen
mit unterschiedlichen funktionellen Gruppen,
Ph.D. Thesis (University of Munich, Munich 2000),
German
32.70 M. Laska: Olfactory discrimination ability of hu-
man subjects for enantiomers with an iso-
propenyl group at the chiral center, Chem. Senses
29, 143152 (2004)
32.71 M. Laska, P. Teubner: Olfactory discrimination
ability of human subjects for ten pairs of enan-
tiomers, Chem. Senses 24, 161170 (1999)
32.72 M. Laska, A. Liesen, P. Teubner: Enantioselectivity
of odor perception in squirrel monkeys and hu-
mans, Am. J. Physiol. 277, R1098R1103 (1999)
32.73 M. Laska, D. Genzel, A. Wieser: The number of
olfactory receptor genes and the relative size of
olfactory brain structures are poor predictors of
olfactory discrimination performance with enan-
tiomers, Chem. Senses 30, 171175 (2005)
Part D | 32
32.74 S. Kim, M. Amundin, M. Laska: Olfactory discrim-
ination ability of South African fur seals (Arc-
tocephalus pusillus) for enantiomers, J. Comp.
Physiol. A 199, 535544 (2013)
32.75 M. Laska, G.M. Shepherd: Olfactory discrimination
ability of CD-1 mice for a large array of enan-
tiomers, Neurosci. 144, 295301 (2007)
32.76 M. Laska, C.G. Galizia: Enantioselectivity of odor
perception in honeybees, Behav. Neurosci. 115,
632639 (2001)
32.77 T. Clarin, S. Sandhu, R. Apfelbach: Odor detection
and odor discrimination in subadult and adult
rats for two enantiomeric odorants supported by
c-fos data, Behav. Brain Res. 206, 229235 (2010)
32.78 B.D. Rubin, L.C. Katz: Spatial coding of enan-
tiomers in the rat olfactory bulb, Nat. Neurosci.
4, 355356 (2001)
32.79 K.J. Rossiter: Structure-odor relationships, Chem.
Rev. 96, 32013240 (1996)
32.80 W.S. Cain: Odor magnitude: Coarse versus fine
grain, Percept. Psychophys. 22, 545549 (1977)
32.81 W.S. Cain: Differential sensitivity for smell: Noise
at the nose, Science 195, 796798 (1977)
32.82 L. Jacquot, J. Hidalgo, G. Brand: Just notice-
able difference in olfaction is related to trigemi-
nal component of odorants, Rhinol. 48, 281284
(2010)
32.83 B.M. Slotnick, J.E. Ptak: Olfactory intensity-dif-
ference thresholds in rats and humans, Physiol.
Behav. 19, 795802 (1977)
32.84 B.M. Slotnick: Olfactory discrimination in rats
with anterior amygdala leasions, Behav. Neurosci.
99, 956963 (1985)
32.85 P. Bonfils, P. Faulcon, L. Tavernier, N.A. Bonfils,
D. Malinvaud: Home accidents associated with
anosmia, Presse Med. 37, 742745 (2008)
32.86 D.V. Santos, E.R. Reiter, L.J. DiNardo,
R.M. Costanzo: Hazardous events associated
with impaired olfactory function, Arch. Otolaryn-
gol. Head Neck Surg. 130, 317319 (2004)
32.87 S. Doucet, R. Soussignan, P. Sagot, B. Schaal: The
smellscape of mothers breast: Effects of odor
masking and selective unmasking on neonatal
arousal, oral, and visual responses, Dev. Psy-
chobiol. 49, 129138 (2007)
32.88 J. Porter, B. Craven, R.M. Khan, S.J. Chang, I. Kang,
B. Judkewicz, J. Volpe, G. Settles, N. Sobel: Mech-
anisms of scent-tracking in humans, Nature Neu-
rosci. 10, 2729 (2007)
32.89 I. Croy, S. Negoias, L. Novakova, B.N. Landis,
T. Hummel: Learning about the functions of the
olfactory system from people without a sense of
smell, PLoS ONE 7, e33365 (2012)
32.90 K. Aschenbrenner, C. Hummel, K. Teszmer,
F. Krone, T. Ishimaru, H.S. Seo, T. Hummel: The
influence of olfactory loss on dietary behaviors,
Laryngoscope 118, 135144 (2008)
32.91 T. Hummel, S. Nordin: Olfactory disorders and
their consequences for quality of life, Acta. Oto-
Laryngol. 125, 116121 (2005)
32.92 C. Moessnang, A. Finkelmeyer, A. Vossen,
F. Schneider, U. Habel: Assessing implicit odor
localization in humans using a cross-modal
spatial cueing paradigm, PLoS ONE 6, e29614
(2012)
32.93 J.D. Porteous: Smellscape, Progr. Phys. Geogr. 9,
356378 (1985)
32.94 M. Beaulieu-Lefebvre, F.C. Schneider, R. Kupers,
M. Ptito: Odor perception and odor awareness in
congenital blindness, Brain Res. Bull. 84, 206
209 (2011)
32.95 R.L. Doty, M.M. Orndorff, J. Leyden, A. Kligman:
Communication of gender from human axillary
odors: Relationship to perceived intensity and
hedonicity, Behav. Biol. 23, 373380 (1978)
32.96 S. Mitro, A.R. Gordon, M.J. Olsson, J.N. Lundstrm:
The smell of age: Perception and discrimination of
body odors of different ages, PLoS ONE 7, e38110
(2012)
32.97 D. Singh, P.M. Bronstad: Female body odour is
a potential cue to ovulation, Proc. Roy. Soc. B 268,
797801 (2001)

32.98 S. Kuukasjrvi, C.J.P. Eriksson, E. Koskela,
T. Mappes, K. Nissinen, M.J. Rantala: Attractive-
ness of womens body odors over the menstrual
cycle: The role of oral contraceptives and receiver
sex, Behav. Ecol. 15, 579584 (2004)
32.99 C.L. Whittle, S. Fakharzadeh, J. Eades, G. Preti:
Human breath odors and their use in diagnosis,
Ann. N.Y. Acad. Sci. 1098, 252266 (2007)
32.100 F. Prugnolle, T. Lefevre, F. Renaud, A.P. Mller,
D. Miss, F. Thomas: Infection and body odours:
evolutionary and medical perspectives, Infect.
Genet. Evol. 9, 10061009 (2009)
32.101 M. Schleidt: Personal odor and nonverbal com-
munication, Ethol. Sociobiol. 1, 225231 (1980)
32.102 R.H. Porter: Olfaction and human kin recognition,
Genetica 104, 259263 (1999)
32.103 G.E. Weisfeld, T. Czilli, K.A. Phillips, J.A. Gall,
C.M. Lichtman: Possible olfaction-ased mecha-
nisms in human kin recognition and inbreeding
avoidance, J. Exp. Child Psychol. 85, 279295
(2003)
32.104 R.H. Porter: Mutual mother-infant recognition in
humans. In: Kin Recognition, ed. by P.G. Hepper
(Cambridge Univ. Press, Cambridge 1991) pp. 413
432
32.105 J. Havliek, P. Lenochova: The effect of meat con-
sumption on body odor attractiveness, Chem.
Senses 31, 747752 (2006)
32.106 J. Albrecht, M. Demmel, V. Schopf, A.M. Klee-
man, R. Kopietz, J. May, T. Schreder, R. Zer-
necke, H. Bruckmann, M. Wiesmann: Smelling
chemosensory signals of males in anxious versus
nonanxious condition increases state anxiety of
female subjects, Chem. Senses 36, 1927 (2011)
32.107 W. Zhou, D. Chen: Entangled chemosensory emo-
tion and identity: Familiarity enhances detection
of chemosensorily encoded emotion, Social Neu-
rosci. 6, 270276 (2011)
32.108 D.L. Wells, P.G. Hepper: The discrimination of dog
odours by humans, Percept. 29, 111115 (2000)
32.109 P.G. Hepper, D.L. Wells: Individually identifiable
body odors are produced by the gorilla and dis-
criminated by humans, Chem. Senses 35, 263268
(2010)
32.110 A.N. Gilbert, K. Yamazaki, G.K. Beauchamp,
L. Thomas: Olfactory discrimination of mouse
Human and Animal Olfactory Capabilities Compared References 689
strains (Mus musculus) and major histocompat-
ibility complex types by humans (Homo sapiens),
J. Comp. Psychol. 100, 262265 (1986)
32.111 J. Havliek, R. Dvoakova, L. Bartos, J. Flegr:
Non-advertized does not mean concealed: Body
odour changes across the human menstrual cycle,
Ethology 112, 8190 (2006)
32.112 C. Ober: Studies of HLA, fertility and mate choice
in a human isolate, Hum. Reprod. Update 5, 103
107 (1999)
32.113 R. Chaix, C. Cao, P. Donnelly: Is mate choice in hu-
mans MHC-dependent?, PLoS Genet. 4, e1000184
(2008)
32.114 J. Havliek, S.C. Roberts: MHC-correlated mate
choice in humans: A review, Psychoneuroen-
docrinology 34, 497512 (2009)
32.115 I. Croy, V. Bojanowski, T. Hummel: Men with-
out a sense of smell exhibit a strongly reduced
number of sexual relationships, women exhibit
reduced partnership security A reanalysis of
previously published data, Biol. Psychol. 92, 292
294 (2013)
32.116 V. Gudziol, S. Wolff-Stephan, K. Aschenbrenner,
P. Joraschky, T. Hummel: Depression resulting
from olfactory dysfunction is associated with re-
duced sexual appetite A cross-sectional cohort
study, J. Sexual Med. 6, 19241929 (2009)
32.117 J.M. Brunstrom: Dietary learning in humans: Di-
rections for future research, Physiol. Behav. 85,
5765 (2005)
32.118 J.A. Gottfried, J. ODoherty, R.J. Dolan: Appeti-
tive and aversive olfactory learning in humans
studied using event-related functional magnetic
resonance imaging, J. Neurosci. 22, 1082910837
(2002)
32.119 D.A. Wilson, R.J. Stevenson: Learning to Smell. Ol-
factory Perception from Neurobiology to Behavior
(Johns Hopkins Univ. Press, Baltimore 2006)
32.120 T. Engen, B.M. Ross: Long-term memory of odors
with and without verbal descriptions, J. Exp. Psy-
Part D | 32
chol. 100, 221227 (1973)
32.121 W.P. Goldman, J.G. Seamon: Very long-term
memory for odors: Retention of odor-name as-
sociations, Am. J. Psychol. 105, 549563 (1992)