C H A P T E R

1
Animal Perception Including
Differences With Humans
Colin G. Scanes
University of Wisconsin–Milwaukee, Milwaukee, WI, United States

1.1 INTRODUCTION paucity of human senses may be viewed as a

Humans and animals use their senses to per-
ceive the environment to locate food, water,
mates, shelter, dangers, such as predators, or to
communicate between other members of their
species. Animals have marked differences in
their sensory systems depending on their evo-
lution and environment. This chapter will con-
sider the senses under the following sections.
• auditory perception in Section 1.2.
• light/visual perception (sight) in Section 1.3.
• chemoreception in Section 1.4.
• tactile perception or touch in Section 1.5.
• other senses in Section 1.6.
These are discussed in detail in succeeding
sections. It should be noted that wild mammals
do not appear to be able to detect many envi-
ronmental stimuli that humans in the developed
societies are able to, by using various artificial
devices. For example, we can detect nonvisual
spectra, such as infrared, radio waves, and gam-
ma radiation using instruments. The relative
detriment or as a means of freeing up neurons
and allowing the brain to be used for other tasks,
such as higher-level thinking. For instance, al-
though Neanderthals and anatomically modern
humans had similarly sized brains, Neander-
thals had larger visual systems and presumably
associated brain areas (Pearce et al., 2013).
1.2  HEARING OR SOUND
DETECTION
Ears detect the auditory frequency of sound
or acoustic waves in the air (or for aquatic mam-
mals in water). Sound varies with wave fre-
quency (pitch) and amplitude (loudness). Low
notes have a relatively low frequency of sound
waves per second (hertz or Hz) while high notes
have a high frequency (high kilohertz or kHz or
Hz × 1000). The ear is made up of the following:
• The external ear consists of the pinnae or
auricles and the ear canals (also named as
the external auditory canals or external

Animals and Human Society

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1 Copyright © 2018 Elsevier Inc. All rights reserved.
2 1.  Animal Perception Including Differences With Humans

auditory meatus). The ear canals end at the
tympanic membrane or eardrum.
• The middle ear with three bones (the
auditory ossicles: malleus, incus, and stapes)
in the air-filled tympanic cavity.
• The inner ear including the semicircular
canals (responsible for balance) and cochlea
(responsible for hearing).
The pinnae have important roles in transmit-
ting sound (Ballachanda, 1997) acting to do the
following:
• filter sound attenuating low-frequency
sound
• amplifying middle frequencies of sound
People and animals can locate the direction
of sounds based on various factors including
the time differences in detection by the two
ears, and amplification of different frequencies
together with movement of the head and body.
In some mammals, the pinnae rotate partially to
aid in location (discussed in Section 1.2.1).
Sound is transmitted to the middle ear; first
inducing vibrations of the tympanic membrane
and then the auditory ossicles in the air-filled
tympanic cavity. The vibrations are detected by
sensory hair cells in the organ of Corti within the
inner ear. This leads to depolarization of the au-
ditory neurons (Uemura, 2015).
1.2.1  Auditory Differences Between
Human and Animals
There are marked differences between the
frequencies of sound detected by the ear in hu-
mans and animals (Table 1.1). Ultrasound is
above the highest range of sound/vibrations of
air normally detected by the human ear (above
20 kHz). Animals including cats, rats, and mice
can detect ultrasonic sound with mice and rats
communicating via ultrasonic vocalization
(Carruthers et al., 2013).
The pinnae can be moved (rotate) in some
animals including cats, mice, rats, some breeds
of dogs, and bats. This facilities detection of
the spatial location of the origin of the sound
(cats, Musicant et al., 1990; Phillips et al., 1982;
reindeer, Flydal et al., 2001). Moreover, this is
used for echolocation in bats with the pinnae
focusing certain frequencies of ultrasonic sound
(Kuc, 2009). The evolution of echolocation is in-
teresting. It is thought that echolocation based
on laryngeal ultrasonic vocalization was devel-
oped in the ancestors of horseshoe bats and Old
World fruit bats. It was subsequently lost in the
ancestors of Old World fruit bats and then re-
gained in some with tongue clicking ultrasonic
vocalization (Jones and Teeling, 2006; Razak
and Fuzessery, 2015; reviewed by Jones and
Holderied, 2007).

TABLE 1.1 Range of Sound Frequencies Detected Between Humans and Different Animal Species
Species Lower limit (Hz) Upper limit (kHz)

Human 20 20
Dog 67 45
Cat 45 78
Mouse 1000 91
Bat 2000 110
Whale 1000 123
Different units in lower and upper limits.
Based on Elert, G., 2003. Frequency range of human hearing. Available from: http://hypertextbook.com/facts/2003/ChrisDAmbrose.shtml; Fay, R.R., 1988.
Hearing in Vertebrates: A Psychophysics Databook. Hill-Fay Associates, Winnetka, IL; Heffner, R.S., Heffner, H.E., 1985. Hearing range of the domestic
cat. Hear. Res. 19, 85–88; Heffner, H.E., Heffner, R.S., 2007. Hearing ranges of laboratory animals. J. Am. Assoc. Lab. Anim. Sci. 46, 11–13; Warfield, D.,
1973. The study of hearing in animals. In: Gay, W. (Ed.), Methods of Animal Experimentation, IV. Academic Press, London, pp. 43–143.

 
 1.3 Vision 3
There are some other differences in the ex-
ternal ear between humans and animals. For
instance, there are no pinnae in birds. Similarly,
there are either no or very small pinnae in marine
mammals of the order Cetacea (dolphins, porpois-
es, and whales) and Pinnipedia (a clade within the
order Carnivora). In Cetaceans, the auditory canal
is not open to the tympanic membrane. Instead
it may be narrow and plugged with wax. In Pin-
nipeds, a valve closes the auditory canal.
It is interesting to also consider hearing in
human ancestors with other great apes. Based FIGURE 1.1  Comparison of what dogs and people see.
on the morphology of the auditory canal, there Source: Based on Coren, S., 2008. Can Dogs See Colors? Psychol-
ogy Today. Available from: http://www.psychologytoday.com/blog/
are differences in sensitivity to frequencies
canine-corner/200810/can-dogs-see-colors.
1.5–3.5 kHz between humans and chimpanzees
and also with the early hominin species, Aus-
tralopithecus africanus and Paranthropus robustus monochromatic light, even dim light, down to a
(Quam et al., 2015). few photons (Solovei et al., 2009). The cones are
responsible for color vision and require bright
light. The photopigment in rods is called rho-
dopsin. In the cones, the photoreceptor mole-
Dog whistles produce a sound in the ul-
cules contain forms of opsin linked to 11-cis reti-
trasonic range above the audible sound fre-
nal. In most mammals but not humans, there are
quency range for humans (>23,000 Hz).
two distinct photoreceptors in the cones detect-
Dogs are used as hearing dogs for the
ing blue and red light (Okano et al., 1992) with
deaf.
the following absorption maxima in dogs (Neitz
et al., 1989):
• blue pigment (absorption maxima 429 nm)
1.3 VISION • red/yellow pigment (absorption maxima
555 nm)
The protein, opsin, is involved in photorecep-
Fig. 1.1 summarizes the approximation of a
tion in all animals. Opsin is an ancient protein
comparison of colors that a dog and a person
found in insects and vertebrates (Yokoyama and
would see.
Yokoyama, 1989) whose common ancestor lived
about 950 million years ago (see Chapter 10,
Section 10.1). In vertebrates at least, the opsin is 1.3.1  Differences in Vision Between
a G protein-linked transmembrane protein. The Humans and Animals
collision of photon(s) with light is detected by
The number of rods and cones are similar at
the interaction of a photon with the 11-cis iso-
least between humans and other primates:
form of retinal and its transformation to an all-
trans confirmation and then induction of confor- • There are 3.1 million cones in rhesus monkey
mation of the opsin molecule. (Wikler et al., 1990) and pigtail macaque
There are two photoreceptor cell types in (Macaca nemestrina) (Packer et al., 1989)
the eye: (1) rods and (2) cones. There are over compared 3.2 million in human (Jonas
20-fold more rods than cones. The rods detect et al., 1992).

 
4 1.  Animal Perception Including Differences With Humans
• There are 61.0 million rods in rhesus monkey
and 60.5 million pigtail macaque (M.
nemestrina) as compared to 60.1 million in
human (Jonas et al., 1992).
However, there are differences between
nocturnal versus diurnal species with larger
eyes and greater density of rods in the retina
of nocturnal species, such as cats, mice, rats,
and deer, compared to diurnal species, such as
cattle and humans (Solovei et al., 2009). More-
over, there are differences in the structure of
the rods (Solovei et al., 2009). Thus, nocturnal
species have much better night vision than di-
urnal species.
There are also differences in cones between
humans and most mammals. In humans, there
are three distinct photoreceptors detecting blue,
green, and red light (Fig. 1.2) (reviewed by Yo-
koyama and Yokoyama, 1989).
• blue pigment (absorption maxima 420 nm)
• green pigment (absorption maxima 530 nm)
• red pigment (absorption maxima 560 nm)
The red and green pigments are found only
within primates and specifically in Old World
monkeys and great apes (clade Catarrhini).
These pigments resulted from gene duplica-
tion about 35–40 million years ago in the com-
mon ancestor of Old World monkeys and great
apes (Ibbotson et al., 1992; Yokoyama and
Yokoyama, 1989). There appears to have been
a parallel development of trichromatic vision
FIGURE 1.2  Comparison of spectra absorption by cone pigments between (A) human and (B) starling. Source: Based on
Osorio, D., Vorobyev, M., 2008. A review of the evolution of animal colour vision and visual communication signals. Vision Res. 48,
2042–2051. Elsevier and open access.
 
and independent gene duplication in the an-
cestors of the New World monkey—the howler
monkey (Dulai et al., 1999). Moreover, the blue
and red/green pigments resulted from a much
earlier gene duplication about 250 million years
ago (Yokoyama and Yokoyama, 1989). In con-
trast to the three photopigments in higher pri-
mates including humans, there are four in chick-
ens and other birds with absorption maxima of
the following: violet/ultraviolet, 415 nm; blue,
455 nm; green, 508 nm; and red, 570 nm (Okano
et al., 1989, 1992) (Fig. 1.2).
1.3.2  Extraretinal Detection of Light
(Pineal and Hypothalamus)
Unlike in humans and other mammals, light
can be detected within the brain, and specifi-
cally the hypothalamus, of birds (reviewed
by García-Fernández et al., 2015). It has been
known since the 1930s that reproductive func-
tioning is influenced by light detected in the hy-
pothalamus (reviewed by Kuenzel et al., 2015).
The opsin, vertebrate ancient opsin, present in
the hypothalamus neuroendocrine cells (e.g.,
in the gonadotropin-releasing hormone ex-
pressing neuroendocrine cells) is different from
that in the rods and cones (reviewed by Davies
et al., 2012). In addition, there is light detection
together with photosensitive proteins expressed
in the avian pineal (Kubo et al., 2006; Okano
et al., 1994).
 1.4  Chemical Senses 5
1.4  CHEMICAL SENSES
Mammals have a least four distinct systems
of chemoreception:
• olfactory chemoreception (sense of smell)
• vomeronasal chemoreception (detecting
pheromones)
• taste or gustatory chemoreception (on
tongue)
• trigeminal chemoreception (for noxious
compounds, e.g., in cheek)
These will be considered next.
1.4.1  Olfactory Chemoreception
Humans and other mammals detect a vast
number of volatile chemicals; these being per-
ceived as odors or odorants. The olfactory sys-
tem consists of the olfactory epithelium lining
the posterior nasal cavity with neural processes
to the olfactory bulb and other brain regions.
The terminals of the olfactory sensory neurons
are present in the olfactory epithelium. These
contain specialized/modified cilia, which con-
tain the olfactory receptors (OR) in the cell mem-
brane. Binding of odorants to specific OR then
leads to the generation of action potentials.
These contain OR. In most mammals, there
are over 1000 different such receptor proteins
encoded by the OR gene family with over 1000
different genes (reviewed by Buck, 2004).
1.4.1.1  Comparison of Olfaction Between
Humans and Animals
Humans have about 350 functioning OR
genes and over 600 OR pseudogenes (Gilad
et al., 2003; reviewed by Buck, 2004). This is
in contrast to the situation in most mammals
where there are over 1000 OR (e.g., mice and
pigs) (Gilad et al., 2003; Nguyen et al., 2012;
reviewed by Buck, 2004) and over 1300 genes
in dogs (Olender et al., 2004). Catarrhinine
primates (Old World monkeys and the great
apes) have increased numbers of OR pseudo-
genes (Gilad et al., 2003; Rouquier et al., 2000)
 
coincident with the acquisition of trichromatic
vision (Gilad et al., 2004). There is evidence that
the human versus the chimpanzee lineage had a
higher rate of loss of OR genes (becoming pseu-
dogenes) (Gilad et al., 2003, 2005). There are also
major losses of the number of functional OR and
the sense of smell in marine mammals (Ceta-
ceans, such as whales and porpoises, and Pin-
niped carnivores, such as sea lions) and reptiles
including sea turtles and sea snakes (Kishida
and Hikida, 2010; Kishida et al., 2007). These are
independent and parallel following the transi-
tion to a marine environment.
Dogs have an exquisite sense of smell due
to the sensitivity of their olfactory system with
over 1300 OR genes. There is evidence for poly-
morphisms in the OR genes with heterozygous
dogs having greater detection abilities (Lesniak
et al., 2008). This olfactory ability is used for
sniffer dogs (see Chapter 2).
1.4.1.2  Odor Detection in Pigeons
It is perhaps surprising that the speed of re-
turn of homing pigeons was reported to be im-
proved under smoggy polluted conditions in
the Peoples Republic of China (Li et al., 2016).
There is evidence that environmental odor plays
a role in homing pigeons (Gagliardo et al., 2011)
and that severing the olfactory nerve impairs
homing (Papi et al., 1971).
1.4.2  Taste or Gustatory
Chemoreception
The ability to taste ingested food and if neces-
sary spit it out is an obvious advantage. If food
or water contains a toxicant, it may be bitter and
not swallowed. A sour taste may reflect spoiled
food. If a food contains sugar and is sweet, it will
be swallowed.
What can we or animals taste? The well-
established tastes in humans and throughout
the vertebrates are the following:
• sweet
• bitter
6 1.  Animal Perception Including Differences With Humans

• sour (acidity or low pH) • The bitter taste receptor is mediated by the
• salty 25 members of the type 2 taste receptor
• umami: savoriness/glutamate, such as in families (Meyerhof et al., 2010).
monosodium glutamate • Salt is detected by epithelial-type sodium
channels leading to depolarization of the
Additional candidate or putative tastes include
receptor cell.
detectors for fatty acids/lipids and water (Bach-
• Sourness is detected by proton (hydrogen
manov and Beauchamp, 2007).
ion) gated cation and chloride channels.
Taste is mediated by taste buds or gustatory
papillae on the surface of the tongue. Specific 1.4.2.1  Is Taste Reception the Same Across
molecular receptors are responsible for taste. Mammals and Birds?
When a specific stimulus binds to the receptor,
In a number of species of the order Carnivora
there is depolarization, which is transmitted to
including cats, there is no taste reception to sug-
the taste receptor cell, and activation of a neuron
ar and other sweeteners. This is due to the gene
[in branches of three cranial nerves: VII (facial),
Tas1r2 becoming inactive as a pseudogene (Jiang
IX (glossopharyngeal), and X (vagus)] via the
et al., 2012; Li et al., 2005, 2009). In an analogous
intervening synapses (reviewed by Bachmanov
manner, there is also loss of functional umami
and Beauchamp, 2007; Purves, 2001). It was as-
taste reception in bats (Zhao et al., 2012). Inter-
sumed that all molecular taste receptors span
estingly, based on genomic data, penguins ap-
the cell membrane. This is true for the taste 1
pear to have lost the ability to taste bitter, sweet,
and 2 families of receptors (T1R and T2R) are
and umami (Zhao et al., 2015). This may be ex-
members of the superfamily of G protein-cou-
plicable in terms of their diets of cephalopods
pled receptors (Zhang et al., 2003; reviewed
and fish or their swallowing food immediately
by Bachmanov and Beauchamp, 2007). These
with no time in the mouth.
consist of extracellular, intracellular, and trans-
membrane domains, the latter across the plasma 1.4.2.2  Presence of Molecular Taste
membrane. These form dimers and activate the Receptors in Other Organs
G protein, gustducin. These receptors are in the
Molecular taste receptors are also present in
apical microvilli of the taste cells (reviewed by
the gastrointestinal tract. For instance, activa-
Bachmanov and Beauchamp, 2007). However,
tion of umami receptors in the colon leads to
some other taste stimuli including sodium, pro-
peristalsis (Kendig et al., 2014) and activation of
tons (pH), some bitter, and some sweet com-
the sweet taste receptors in the small intestine
pounds enter the cell.
leads to the release of glucagon-like peptide 1
Sweet taste is primarily mediated by a het-
(Jang et al., 2007) and the expression of sodium
erodimer of two members of the type 1 taste
glucose cotransporters (Margolskee et al., 2007).
receptor family, Tas1r2/Tas1r3; the heterodimer
The impact of gastrointestinal molecular taste
being two different molecules linked together as
receptors in livestock or poultry has received
a functional unit (reviewed by Bachmanov and
little attention.
Beauchamp, 2007). The molecular taste recep-
tors are the following:
• The principal umami taste receptor is 1.4.3  Vomeronasal Chemoreception
a heterodimer of two members of the The vomeronasal or Jacobson’s organs (VNO)
type 1 taste receptor family, Tas1r1/ detect pheromones and other chemical signals
Tas1r3 (reviewed by Bachmanov and by their binding to VNO chemoreceptor cells.
Beauchamp, 2007). [It should be noted that pheromones influence

 
 1.5 Touch 7
the VNO and olfactory epithelium with the re-
sponse in pigs to the pheromone androstenone
not mediated via the VNO (Dorries et al., 1997).]
Activation of the VNO by specific pheromones
leads to physiological or behavioral changes
(reviewed by Dulac and Axel, 1995). There are
two gene families of VNO chemoreceptors,
V1R and V2R, with over 100 genes (Dulac and
Axel, 1995). The V1R and V2R proteins have
seven transmembrane domains and are linked
to G proteins. The activation of the V1R or V2R
proteins increases intracellular inositol 1,4,5-tri-
sphosphate and hence with an ion channel of the
transient receptor potential family (TRP) playing
a critical role in signal transduction (reviewed
by Keverne, 1999; Zhang and Webb, 2003).
The tubular VNO are found at the base of the
nasal septum or in the roof of the mouth. The
pheromones are detected by chemosensory neu-
rons with axonal projections to the accessory
olfactory bulb and other centers, such as in the
amygdala and hypothalamus (reviewed by Du-
lac and Axel, 1995).
1.4.3.1  Differences Between Humans and
Animals: Vomeronasal Chemoreception
There is still controversy as to whether or
not adult humans even have vomeronasal or-
gans and whether humans have functional
vomeronasal chemoreceptors (reviewed by
Meredith, 2001). Perhaps this is not surprising
given that the existence of human pheromones
is still not fully established (reviewed by Mere-
dith, 2001). Similarly, there appears to be loss of
vomeronasal signaling in bats (Zhao et al., 2011).
Components of the VNO chemoreception sys-
tem are not functional throughout the catarrhine
primates (Old World monkeys and great apes)
with V1R and TRP genes becoming nonfunc-
tional or pseudogenes (Zhang and Webb, 2003).
It has been suggested that the reddening and
swelling of the vulva at the time of estrus in Old
World monkeys is a superior signal than phero-
mones and made the VNO dispensable (Zhang
and Webb, 2003). Alternatively, the ability to
 
discern differences between reds, oranges, and
greens would have distinct advantages for
fruit-eating animals during foraging (Surridge
et al., 2003).
1.4.4  Trigeminal Chemoreception
Trigeminal nerve terminals can be activated
by capsaicin, a chemical responsible for the sen-
sation of spicy hot foods containing chili pep-
pers or plants of the genus Capsicum (Liu and
Simon, 2000). In addition, the trigeminal nerve
terminals detect acid (e.g., in vinegar), acting
via H+-gated ion channels, and other noxious
stimuli. These include air pollutants (e.g., sulfur
dioxide and ammonia) and carbon dioxide (e.g.,
in cold sodas).
1.4.4.1  Animal–Human Differences in
Trigeminal Responses
The trigeminal system of birds is not sensitive
to capsaicin, the usual model for trigeminal che-
moreception in mammals (Mason et al., 1991).
1.5 TOUCH
1.5.1  Animal–Human Differences: People
Do Not Have Whiskers or Vibrissae
Vibrissae are stiff specialized hairs. They
function as tactile or touch sensory organs. They
can be either macrovibrissae (long whiskers)
or microvibrissae (Prescott et al., 2014). Macro-
vibrissae are found on the face of most mam-
mals including rodents, carnivores, and many
primates (Fig. 1.3). Humans are one of the few
mammals that are devoid of vibrissae. It has
been suggested that vibrissae are particularly
useful to provide an awareness of the environ-
ment in nocturnal species.
The movement of or shear forces in vibrissae
are transduced to Merkel cells in the epidermis
and then via a synapse-like process to adren-
ergic afferent neurons (Cha et al., 2011; Ikeda
8 1.  Animal Perception Including Differences With Humans

FIGURE 1.3  Vibrissae are sense organs in various animals, but have been lost in humans through evolution. (Top row)
Vibrissae by the nose of a dog. (Middle row) Vibrissae by the nose and mouth of a cat and rats. (Bottom row) Vibrissae on sea
lion and walrus. Source: From Wikimedia Commons.

et al., 2014; Ma, 2014; Maksimovic et al., 2014).
In rodents, vibration of the vibrissae plays a
role in intraspecies communication (Mitchinson
et al., 2011).
Based on a comparison of the human ge-
nome to that of the chimpanzee and other spe-
cies, it was possible to identify deletions in the
human genome responsible for the loss of the
vibrissae enhancer from the human androgen
receptor (McLean et al., 2011). There was also
a loss of penile spines as a result of this dele-
tion (McLean et al., 2011) leading to desensiti-
zation of the penis, longer duration of coitus,
and presumably strengthening of bonding (van
Driel, 2011). While people do not have vibrissae,
cancers of the Merkel cells is an uncommon but
highly aggressive skin carcinoma in people (Sib-
ley et al., 1980; Tilling and Moll, 2012) and dogs
(Joiner et al., 2010).
1.6  OTHER SENSES
Birds have two magnetoreceptor systems:
one based on magnetite in or near the beak, and
the other in the retina based on putative magn-
etosensory molecules, the cryptochromes. These
allow detection of the direction of the Poles and

 
 REFERENCES 9
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Further Reading
Douglas, R.H., Jeffery, G., 2014. The spectral transmission
of ocular media suggests ultraviolet sensitivity is wide-
spread among mammals. Proc. Biol. Sci. 281, 20132995.
Pinc, L., Bartoš, L., Reslová, A., Kotrba, R., 2011. Dogs dis-
criminate identical twins. PLoS One 6, e20704.