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Olfaction
Chapter 13

Olfaction

Olfactory Physiology From Chemicals to Smells Olfactory Psychophysics, Identification, and Adaptation Olfactory Hedonics Olfaction, Memory, and Emotion

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Odors: Olfactory sensations Chemical compounds.

Olfactory Physiology

But not every chemical is odorant. In order to be smelled: Molecule must be volatile, small, and hydrophobic.

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Odorants

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The human olfactory apparatus:

Olfactory Physiology

Primary purpose of nose: to filter, warm, humidify air we breathe. Nose: Small ridges, olfactory cleft, olfactory epithelium.

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The Nose

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Olfactory Physiology

The olfactory epithelium: the retina of the nose: Three types of cells: 1. Supporting cells 2. Basal cells 3. Olfactory sensory neurons (OSNs)cilia protruding into mucus covering olfactory epithelium

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Olfactory Physiology

The olfactory epithelium: the retina of the nose: Olfactory receptors (ORs): Interaction between these and odorant stimulates cascade of biochemical events

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Olfactory Sensory Neuron

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Olfactory Perception Pathway

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Sense of taste. Danger warning.

Olfactory Physiology

Olfactory loss can cause great suffering:

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Olfactory Physiology

Olfactory receptor cells are different from all other sensory receptor cells: They are not mediated by any protective barrier, make direct contact with brain. (e.g., visual receptors are protected by cornea, receptors for hearing are protected by eardrum, taste buds are buried in papillae). Therefore many drugs can be inhaled.

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Olfactory Physiology

Olfactory receptor cells are different from all other sensory receptor cells: They are not mediated by any protective barrier, make direct contact with brain. OSN axons are among thinnest and slowest in body. Therefore it takes a long time to perceive odors compared to other perceptions.

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Olfactory Physiology

Subtle differences between sensation and perception: Sensation occurs when scent is neurally registered. Perception occurs when becoming aware of detecting it.

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Olfactory Physiology

The genetic basis of olfactory receptors: In 1991 Buck and Axel showed that genome contains about 1000 different olfactory receptor genes, each codes for single type of OR. Animals have less ORs. Trade-off between vision and olfaction.

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The feel of scent:

Olfactory Physiology

Odorants can stimulate somatosensory system through polymodal nociceptors (touch, pain, temperature receptors). These sensations are mediated by the trigeminal nerve (cranial nerve V). Often, it is impossible to distinguish between sensations traveling up cranial nerve I from olfactory receptors and those traveling up cranial nerve V from somatosensory receptors.

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From Chemicals to Smells

Theories of olfactory perception: Shape-pattern theory: Match between shapes of odorants and odor receptors; dominant biochemical theory. Recent molecular research: Scents are detected by means of combinatorial codes. Vibration theory: There is a different vibrational frequency for every perceived smell, molecules that produce same vibrational frequencies will produce same smell.

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Presumed Odorant-Receptor Binding

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Specific anosmia:

From Chemicals to Smells

The inability to smell one specific compound with otherwise normal smell perception. 50% of population has specific anosmia to androstenone. Change in detection can occur with training. This cannot be explained by vibration theory.

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Study of stereoisomers:

From Chemicals to Smells

Molecules that are mirror-image rotations of one another; although they contain the same atoms, they can smell completely different. Vibration theory cannot explain this phenomenon.

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Stereoisomers

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The importance of patterns:

From Chemicals to Smells

How can we detect so many different scents if our genes only code for about 1000 olfactory receptors? We can detect the pattern of activity across various receptor types. Intensity of odorant also changes which receptors will be activated.

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The importance of patterns:

From Chemicals to Smells

So weak concentrations of odorant do not smell the same as strong concentrations of it. Specific time order is important.

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Hypothetical Role of OR Receptor Activation

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Odor mixtures:

From Chemicals to Smells

We rarely smell pure odorants, rather we smell mixtures. How do we process the components in an odorant mixtures. Two possibilities: 1. Analysis 2. Synthesis

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From Chemicals to Smells

Analysis: Example from auditory mixtures: high note and low note. Synthesis: Example from color mixtures: mixing red and green lights, resulting in yellow light.

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Synthesis

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Odor imagery:

From Chemicals to Smells

Humans have little or no ability to conjure odor images. We do not think in smell, do not imagine smells.

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Olfactory Psychophysics, Identification, and Adaptation

Detection, discrimination, and recognition How much stimulation is required before we perceive something to be there? Olfactory detection thresholds: Depend on several factors.

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Olfactory Psychophysics, Identification, and Adaptation

Women: Generally lower thresholds than men, especially during ovulatory period of menstrual cycles, but their sensitivity is not heightened during pregnancy. Professionals can distinguish up to 100,000 odors (e.g., professional perfumers, wine tasters). Durability: Our recognition of smells is durable even after several days, month, or year.

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Odor Recognition Retention

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Olfactory Psychophysics, Identification, and Adaptation

Identification: Attaching verbal label to smell is not easy. Tip-of-the-nose phenomenon. Compare to tip-of-the-tongue phenomenon. Anthropologists found that there are fewer words for experience of smells as opposed to other sensations.

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Olfactory Psychophysics, Identification, and Adaptation

Sense of smell and language: Disconnected, possibly because: Olfactory information is not integrated in thalamus prior to processing in cortex Majority of olfactory processing occurs in right side of brain while language processing occurs in left side of brain

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Description of Colors vs. Odors

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Olfactory Psychophysics, Identification, and Adaptation

Adaptation: Sense of smell is essentially a change detector. Examples: Walking into bakery, smelling strong perfume that person cannot smell. Receptor adaptation. Cross adaptation.

13 Pathway for Odorant Molecule-Odorant Receptor Binding (1)

13 Pathway for Odorant Molecule-Odorant Receptor Binding (2)

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Olfactory Psychophysics, Identification, and Adaptation

Cognitive habituation After long-term exposure to an odorant, one is no longer able to detect that odorant or has very diminished detection ability. Example: Going out of town, coming back to house with smell.

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Olfactory Psychophysics, Identification, and Adaptation

Three mechanisms involved in cognitive habituation: 1. Olfactory receptors internalized into cell bodies during odor adaptation may be hindered after continuous exposure, take longer to recycle. 2. Odorant molecules may be absorbed into bloodstream causing adaptation to continue. 3. Cognitive-emotional factors.

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Olfactory Psychophysics, Identification, and Adaptation

The importance of attention We cannot smell while we are asleep. Attention is cut off during sleep, therefore we cannot smell.

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Odor hedonics:

Olfactory Hedonics

The liking dimension of odor perception; typically measured with scales pertaining to an odorants perceived pleasantness, familiarity, and intensity.

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Familiarity and intensity:

Olfactory Hedonics

We tend to like odors weve smelled many times before. Intensity: Interesting relationship to odor liking.

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Odorant Pleasantness vs. Intensity

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Nature or nurture?

Olfactory Hedonics

Are hedonic responses to odors innate or learned? Debate over this. Evidence from infants. Cross-cultural data supporting associative learning. An evolutionary argument. Learned taste aversions.

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Olfactory Hedonics

Two caveats for theory that odor hedonics are mostly learned: Trigeminally irritating odors may elicit pain responses, and all humans have innate drive to avoid pain. There is potential variability in receptor genes and pseudogenes that are expressed across individuals.

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Olfaction, Memory, and Emotion

Are odors really the best cues to memory? Other modalities can elicit memory as well (e.g., vision, touch, taste). Memories triggered by odor cues are distinctive in their emotionality. Emotion and evocativeness of odor-elicited memories lead to false impression that such memories are especially accurate.

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Memory Cues

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Orbitofrontal cortex:

Olfaction, Memory, and Emotion

Olfaction is processed there. Also the cortical area for assigning affective value, (i.e., hedonic judgment).