Obial, Erika Kate

Wong, Michelle

I. Intro
A. Unique properties of chemical senses
a. Involve taking molecules into the body
b. Receptors that serve taste and smell undergo a cycle of birth, development, and death
(also knowns as neurogenesis) over 5-7 weeks for olfactory receptors and 1-2 weeks for
taste receptors
B. Chemical senses as molecule detectors - endow gas and liquid molecules with distinctive smell
and tastes
C. Sense of taste and smell as “gate-keepers” of the body
a. Identify things that the body needs for survival
b. Detect things that would be bad for the body and should therefore be rejected
c. Gatekeeper function of taste and smell is aided by a large affective, or emotional,
component
i. Things that are bad for us often taste or smell unpleasant, things that are good
for us generally taste or smell good
ii. Smelling an odor associated with a past place or event can trigger memories
which in turn may create emotional reactions.
II. Taste
A. Taste occurs when molecules enter the mouth in solid or liquid form and stimulate taste
receptors on the tongue
B. Perception resulting from this stimulation have been described in five basic taste qualities
a. Five basic taste sensations: salty, sour, sweet, bitter, and umami (meaty, brothy, or
savory often associated with the flavor-enhancing properties of MSG)
b. Donald McBurney
i. Found that some substances have a predominant taste and that other
substances result in combinations of four tastes
C. Taste accomplishes its gatekeeper function by the connection between taste quality and a
substance’s effect
a. Sweet compounds
i. Often associated with compounds that have nutritive or caloric value, therefore,
important for sustaining life
ii. Cause an automatic acceptance response
iii. Trigger anticipatory metabolic responses that prepare the gastrointestinal
system for processing these substances
b. Bitter compounds
i. Trigger automatic rejection responses to help the organism avoid harmful
substances (eg. poisons such as arsenic and cyanide)
c. Salty tastes
i. Often indicate presence of sodium
ii. When people are deprived of sodium or lose a great deal of sodium through
sweating, they often seek out foods that taste salty in order to replenish the salt
their body needs
D. Neural Code for Taste Quality
a. Structure of the taste system
i. Process of tasting begins with the tongue
 ii. Surface of the tongue contains many ridges and valleys caused by the presence
of structures called papillae, which fall into four categories:
1. Filiform papillae - shaped like cones and are found over the entire
surface of the tongue, giving its rough appearance
2. Fungiform papillae - shaped like mushrooms and are found at the tip
and sides of the tongue
3. Foliate papillae - series of folds along the back of the tongue on the
sides
4. Circumvallate papillae - shaped like flat mounds surrounded by a trench
and are found at the back of the tongue
iii. All of the papillae except the filiform contain taste buds and the whole tongue
contains about 10, 000 buds
iv. Each taste bud contains 50 to 100 taste cells, which have tips that protrude into
the taste pore.
v. Transduction occurs when chemicals contact receptor sites located on the tips
of these taste cells. Electrical signals generated in the taste cells are transmitted
from the tongue in a number of different nerves
1. Chorda tympani nerve (from taste cells on the sides of the tongue)
2. Glossopharyngeal nerve (from the back of the tongue)
3. Vagus nerve (from the mouth and throat)
4. Superficial petrosal nerve (from the soft palette--the top of the mouth)
The fibers from the tongue, mouth, and throat make connections in the brain
stem in the nucleus of the solitary tract. From there, the signals travel to the thalamus
and then to two areas in the frontal lobe that are considered to be the primary cortex--
the insula and the frontal operculum--which are partially hidden behind the temporal
lobe
b. Population coding - the idea that quality is signaled by the pattern of activity distributed
across many neurons
i. Robert Erickson experiment
1. Presented a number of different taste stimuli to a rat’s tongue and
recorded the response of the chorda tympani nerve
2. Hypothesis: of rat’s perception of taste depends on population coding,
two substances with similar patterns should taste similar.
ii. Susan Schiffman and Robert Erickson experiment
1. Asked humans to make similarity judgments between a number of
different solutions
2. Found that solutions judged more similar psychophysically had similar
patterns of firing, as population coding would predict.
c. Specificity coding - the idea that quality is signaled by the activity in individual neurons
that are tuned to respond to specific qualities
Evidence:
i. Ken Mueller and coworkers experiment
1. Used a chemical compound called PTc that tastes bitter to humans but
is not bitter to mice. A specific receptor had been identified as being
responsible for the bitter taste of PTC in humans.
2. Used genetic cloning techniques to create a strain of mice that had this
human bitter-PTC receptor. Mice with this receptor avoided high
concentrations of PTC.
 3. In another experiment, created a strain of mice that lacked a bitter that
responds to a compound called Cyx. The mice lacking this receptor did
not avoid Cyx and Cyx no longer caused any firing in nerves receiving
signals from the tongue. Therefore, when taste receptor for a substance
is removed, this is reflected in both nerve firing and animal’s behavior.
Note: adding or eliminating bitter receptors had no effect on neural firing and
behavior to sweet, salty, or umami stimuli.
ii. Single neurons respond to taste stimuli
1. Recordings from neurons at the beginning of the taste systems of
animals (ranging from rats to monkeys) revealed that neurons are
specialized to respond to specific stimuli, as well as neurons that
respond to a number of different types of stimuli
iii. Effect of substance amiloride, which blocks the flow of sodium into taste
receptors.
1. Applying amiloride to the tongue affects the salt taste on foods
iv. David Smith and Thomas Scott
1. Argue for population coding based on the finding that at more central
locations in the taste system, neurons are tuned broadly, with many
neurons responding to more than one taste quality.
v. Smith and coworkers
1. Point out that just because there are neurons that respond best to one
compound like salty or sour, this doesn’t mean that these tastes are
signaled by just one type of neuron.
vi. Basic taste qualities might be determined by a specific code but population
coding could determine subtle differences between tastes within a category →
explain why not all substances in a particular category have the same taste (eg.
taste of all sweet substances is not identical)
E. Individual Differences in Taste
a. There are genetic differences that affect people’s ability to sense the taste of certain
substances (eg. people’s ability to taste the bitter substance PTC)
b. What causes these differences?
i. Number of taste buds
ii. Specialized receptors
III. Olfaction and Flavor
A. Functions of olfaction
a. Provide information that can be important for survival
i. Alarm system that alerts us to spoiled food, leaking gas, or smoke from a fire
ii. Olfaction is the primary window to the environment of many species
1. Macrosmatic - having a keen sense of smell that is important to their
survival (eg. many animals)
a. Olfaction provides cues to orient them in space, to mark
territory, and to guide them to specific places, other animals,
and food sources.
b. Olfaction is also extremely important in sexual reproduction
because it triggers mating behavior in many species
2. Microsmatic - having a less keen sense of smell that is not crucial to
their survival (eg. humans)
 iii. Isolated congenital anosmia (ICA) - a condition in which people are born with a
lifelong inability to smell
b. Enriches our lives by combining with taste to create flavor
B. Olfactory Abilities
a. Detecting odors
i. Detection threshold for odors - the lowest concentration at which an odorant
can be detected
ii. Humans are much less sensitive to odors than many animals
1. Rats are 8 to 50 times more sensitive to odors than human, and dogs
are from 300 to 10,000 times more sensitive, depending on the odorant
2. Humans’ individual olfactory receptors are as sensitive as any animal’s.
However, humans are less sensitive to odors because humans have far
fewer receptors than animals.
b. Discriminating between odors
i. Odor discrimination - telling the difference between different odors
ii. One of the challenges of odor discrimination is the cast numbers of odors that
are possible, because most natural olfactory stimuli are mixtures of a large
number of components
iii. Caroline Bushdid and coworkers
1. Proposed that humans can discriminate the difference in the smells of
more than 1 trillion olfactory stimuli.
c. Identifying odors
i. Humans find it difficult to accurately identify specific odors
ii. J. A. Desor and Gary Beauchamp experiment
1. Found that when they presented participants with the names of the
substances at the beginning of the experiment and then reminded them
of the correct names when they failed to respond correctly on
subsequent trials, they could, after some practice, correctly identify 98%
of the substances.
iii. Difficulty in identifying odors results not from a deficiency in our olfactory
system, but from an inability to retrieve the odor’s name from our memory
d. Individual differences in olfaction
i. Genetic differences
1. Sensitivity to chemical β-ionone
2. Smell of steroid
C. Analyzing Odorants: The Mucosa and Olfactory Bulb
a. The Puzzle of Olfactory Quality
i. Creating a way to organize odors and to relate odors to physical properties of
molecules has proven extremely difficult
1. Lack of specific language for odor quality
2. Some molecules that have similar structures can smell different and
molecules that have very different structures can smell similar
3. Odors we encounter everyday consist of mixtures of many chemicals
ii. Odor objects – sources of odors
iii. Perceiving odor objects involves olfactory processing that occurs in two stages:
1. First stage takes place in the olfactory mucosa and olfactory bulb, and
involves analyzing. Olfactory system analyzes the different chemical
 components of odors and transforms these components into neural
activity at specific places in the olfactory bulb.
2. Second stage takes place in the olfactory cortex and beyond and
involves synthesizing. Olfactory system synthesizes information about
chemical components received from the olfactory bulb into
representations of odor objects.
*involves learning and memory
b. The Olfactory Mucosa
i. A dime-sized region located on the roof of the nasal cavity just below the
olfactory bulb
ii. Contains olfactory receptor neurons (ORNs), which are dotted with molecules
called olfactory receptors that are sensitive to chemical odorants. A particular
ORN contains only one type of olfactory receptor. There are about 400 different
types of olfactory receptors, each sensitive to a particular group of odorants.
iii. Odorant molecules are carried into the nose in an air stream, which brings these
molecules into contact with the mucosa
c. How Olfactory Receptor Neurons Respond to Odorants
i. Bettina Malnic and coworkers experiment
1. Recognition profile - pattern of activation for each odorant
2. Each odorant causes a different pattern of firing across ORNs. Odorants
that have similar structures often have similar profiles. However, this
doesn’t always occur.
d. The Search for Order in the Olfactory Bulb
i. Activation of receptors in the mucosa causes electrical signals in the ORNs that
are distributed across the mucosa. These ORNs send signals to glomeruli in the
olfactory bulb
1. Each type of ORN send their signals to just one or two glomeruli
ii. Naoshige Uchida and coworkers & 2-deoxyglucose technique
1. Results show that different odorants cause distinctive patterns of
activation. These also support the idea that there is a map of odorants
in the olfactory bulb called chemotopic map (based on molecular
features of odorants such as carbon chain length or functional groups)
D. Representing Odors in the Cortex
- Where are signals transmitted when they leave the olfactory bulb?
- Olfactory mucosa -> [ Olfactory bulb <-> Piriform Cortex <-> Orbitofrontal Cortex ] <->
Amygdala
- Two main olfactory areas: Piriform Cortex (Primary) and Orbitofrontal
Cortex(Secondary)
a. How odorants are represented in the Piriform Cortex
i. Recall
1. Odors that smell different causes different patterns of firing of olfactory
receptors.
2. Moving to the olfactory bulb, different chemicals cause activity in
specific areas which leads to the proposed odotopic maps
ii. At the Piriform Cortex (PC) the odotopic map vanishes
1. Odorants that caused activity in specific locations in the olfactory bulb
causes widespread activity in the PC, showing an overlap between
activity caused by different odorants
 2. Study by B.F Osmanski and Coworkers (2014)
a. Determined brain activity by measuring changes in blood flow
using functional ultrasound imagery (like fMRI)
b. Showed that the chemicals hexanal and pentyl acetate caused
the activity in the entire PC
3. Study by Robert Rennaker and Coworkers (2007)
a. Used multiple electrodes to measure neural responding in the
PC
b. Isoamyl acetate causes activation across cortex
b. How Odor Objects are Represented
i. How is the olfactory system able to determine the identity of a mystery odor
based on the information in a first-time response?
1. It was proposed that formation of odor objects involves learning, which
links the scattered activations that occur for an object
2. Signals from the olfactory bulb are transformed into a scattered pattern
of activation in the PC
ii. What happens at a First Time Response (i.e. a new flower’s odor)
1. Activated neurons aren’t associated with each other yet at the first
response
2. After a number of exposures to the flower, causing the same activation
pattern to occur over and over, neural connection form and neurons
become associated with each other
3. Pattern of activation has been created that represents the odor
iii. Learning plays an important role in perceiving odors
1. Study by Donald Wilson (2003)
a. Measured response of neurons in a rat’s PC to 2 odorants: (1)
mix of isoamyl acetate (banana + peppermint smell), (2)
component isoamyl acetate alone
b. Interested to see if the neurons can tell the difference
c. Procedure: they presented the mix to the rather for either a
brief exposure (10 secs/ 20 sniffs) or longer exposure (50
sec/100 sniffs) and, after a pause, measured the response to the
mixture and component
d. Results: neurons were able to tell the difference between the
mixture and the component after the 2nd set up (100 sniffs)
e. Take away: given enough time, neurons in the PC can learn to
discriminate between different odors
iv. Not all odor objects require learning
1. I.e.pheromones
a. Pheromone responses may be determined by a second pathway
for olfactory perception that sends its signals and does not
depend on experience for identifying odors
b. Also known as the Dual Pathway Idea
i. Odor objects -> PC
ii. Pheromones/Innate Chemicals -> separate pathway
that creates automatic responses to specific odors
IV. Perception of Flavor
- Taste or the experience of taste is usually a combination of taste, where there is stimulation of
the receptors in the tongue, and olfaction, which is the stimulation of the receptors in the
olfactory mucosa
- The combination of taste and olfaction is called flavor
- Flavor is defined as the overall impression we experience from the combination of nasal and
oral stimulation (Lawless, 2001; Shepherd, 2012) and the perceptions such as the texture and
temperature of food, color of food, and sounds of “noisy” food
- Smell Affects Flavor: “tasting with/without the noise”
- The quality and the intensity of the taste of food changes when you pinch your nostrils,
or even one nostril, shut
- As flavor is the combination of taste and smell, holding your nose changes the flavor as
you mess with the smell or eliminate the olfactory component all together
- The interaction between taste and olfaction occurs at two levels: first in the mouth and nose,
and then the cortex
A. Taste and Olfaction Meet in the Mouth and Noise
a. Chemicals in what we eat cause taste when they activate taste receptors on the tongue
b. Food and drink also release volatile chemicals that reach the olfactory mucosa by
following the retronasal route, which is the passage that connects the oral and nasal
cavities
i. Mouth -> nasal pharynx
ii. Pinching ones nostrils does not close the nasal pharynx but prevents vapors
from reaching the olfactory receptors by eliminating the circulation of air
through the channel
c. Localization of Flavor
i. Oral Capture- Created when food and drink stimulate tactile receptors in the
mouth
1. When you taste food, you experience flavor, and the fact that it is all
happening in your mouth is an illusion created by oral capture (Todrank
& Bartoshuk 1991)
ii. Solutions are more difficult to identify when the nostrils are pinched shut and
are often judged to be tasteless (Mozell, et al 1969)
1. Sodium oleate (strong soapy flavor) and Ferrous sulfate (metallic flavor)
yet tasteless when nostrils are closed
iii. Compounds are not influenced by olfaction
1. Monosodium glutamate (MSG) has the same flavor regardless, making
the sense of taste predominate
B. Taste and Olfaction Meet in the Nervous System
a. Our perceptual experience of their combination is created when they interact in the
cortex
i. Olfactory Pathway from Figure 15.17b with the taste pathway added shows the
connections between olfaction and taste
ii. Vision and touch sends signals to the amygdala (vision), structures in the taste
pathway (touch) and the orbitofrontal cortex (vision and touch)
iii. Interactions of these senses show the multimodal nature of our experience of
flavor
1. The orbitofrontal cortex contains bimodal neurons which are neurons
that respond to more than one sense (i.e. taste and smell or taste and
vision)
 a. Bimodal neurons often respond to similar qualities (i.e. neurons
that responds to the taste of sweet fruits will respond to the
smell of sweet fruits)
b. Orbitofrontal cortex is a cortical center for detecting flavor and
for the perceptual representation of food (Rolls & Baylis, 1994)
c. The insula, the primary taste cortex, is involved in the
perception of flavor
iv. Flavor isn’t a fixed response automatically determined by chemical properties
of food
1. Chemicals in particular food may activate the same pattern of ORNs in
the mucosa but they can be affected by cognitive factors and the amt of
food consumed
C. Flavor is Influenced by Cognitive Factors
a. Expectations can influence both the experience and neural responding of eating food
i. Hilke Plassmann and Coworkers (2008)
1. Subjects in a brain scanner judged the “taste pleasantness” of different
samples of wine
a. They were asked to indicate how much they likes 5 different
wines identified by their price, although 2 wines were repeated
twice
b. Results: wines with no labels are rated the same but when
preceded by the price, the higher priced wine gets a much
higher rating than the lower priced one even when they were
the same wine
c. Also found that labels influenced the response of the
orbitofrontal cortex, with the higher priced wine getting a much
larger response
ii. De Araujo et al (2005)
1. Subjects rated the same odor as more pleasant when labeled “cheddar
cheese” than when it was called “body odor” with the orbitofrontal
cortex responding largely to cheddar cheese label
D. Flavor is influenced by Food Intake: Sensory-Specific Satiety
a. Satiety- when one is full or don’t want to eat anymore
b. Food consumed to satiety is often considered less pleasurable than food consumed
when hungry
c. John O’Doherty and Coworkers (2000)
i. Study showed that both the pleasantness of a food-related odor and the brain’s
response to the odor can be influenced by satiety
1. Subjects underwent two conditions: (1) when hungry, (2) after eating
bananas until satiety, and were under a brain scanner when asked to
judge the pleasantness of the smell of banana and vanilla
a. Pleasantness was the same under condition 1, but decreased for
both smells (vanilla by a little but still positive, and banana by a
lot and became negative) after eating to satiety
d. Sensory-specific Satiety is when it is found that there is a larger effect on the odor
associated with the food eaten to satiety which also occurs in the response of the
orbitofrontal cortex
i. Also occurs in the Insula and Amygdala but not for all subjects
 e. Orbitofrontal cortex may be involved in determining the reward value of foods
i. Food is more rewarding when hungry than as it is consumed
ii. When one reaches satiety, the reward is gone and eating stops
V. Application/Importance
A. Importance of the orbitofrontal cortex and Chemical senses
a. Orbitofrontal cortex sends signal to the hypothalamus (responsible for sight, taste, and
smell of food when hunger is present)
i. Chemical senses processed are there to help guide behavior –avoiding harmful
substances, seeking out nutrients, and helping control the amount of food
consumed (satiety)
ii. i.e. situation: when you’re so full that the smell of food makes you want to
vomit
B. Packaging and Consumer Psychology
a. Tricking the senses to make food taste better or look more appealing to the market
i. Charles Spence in the Accounting for Taste reading
C. Proust Effect: Memories, Emotions, and Smells
a. Taste and olfaction, or flavors, can unlock memories or remind people of events past
b. Research shows a link between odors and specific aspects of memory
i. Rachel Herz and Jonathan Schooler (2002)
1. Had subjects describe a personal memory associated with objects and
then were presented with an object in either visual or odor form
2. Subjects who smelled the odor rated their memories as more emotional
and feeling stronger feelings of “being brought back” to that memory
D. The Impossible Burger
a. Link: https://www.youtube.com/watch?v=R_1VRJAuTy4