CH 13 Olfaction Definitions Olfaction sense of smell Gestation sense of taste Odor general smell sensation of a particular quality, e.g.

e.g. cake had a chocolate odor; by contrast, when referring to specific chemical entity, term odorant should be used Odorant any specific aromatic chemical, e.g. you were given the odorant menthol to smell Olfactory cleft narrow space at back of nose into which air flows, where the main olfactory epithelium is located Olfactory epithelium secretory mucosa in human nose whose primary function is to detect odorants in the inspired air; located on both sides of the upper portion of the nasal cavity and the olfactory clefts, the olfactory epithelium contains 3 types of cells: olfactory sensory neurons, basal cells, and supporting cells Supporting cells one of 3 types of cells in olfactory epithelium; cell type provides metabolic and physical support for the OSNs Basal cells one of 3 types of cell sin olfactory epithelium; precursor ells to olfactory sensory neurons Olfactory sensory neurons (OSNs) main cell type in olfactory epithelium; OSNS are small neurons located beneath a watery mucous layer in epithelium; cilia of OSN dendrites contain receptor sites for odorant molecules Cilia hairlike protrusions on the dendrites of OSNs; receptor sites for odorant molecules are on the cilia, which are the first structures involved in olfactory signal transduction Olfactory receptor (OR) region on cilia of OSNs where odorant molecules bind Cribriform plate bony structure riddled w/ tiny holes at the level of the eyebrows that separates the nose from the brain; the axons from the OSNs pass through the tiny holes of the cirbriform plate to enter the brain Anosmia total inability to smell, most often resulting from sinus illness or head trauma Olfactory (I) nerves first pair of cranial nerves; axons of OSNs bundle together after passing thru cribriform plate to form the olfactory nerve Olfactory bulb blueberry-sized extension of brain just above nose, where olfactoryinfo is first processed; there are 2 olfactory bulbs, 1 in each hemisphere, corresponding to R and L nostrils Ipsilateral same side of body/brain Mitral cells main projective output neurons in olfactory bulbs Tufted cells secondary class of output neurons in the olfactory bulbs Glomeruli spherical conglomerates containing the incoming axons of the OSNs. Each OSN converges onto 2 glomeruli (one medial, one lateral) Primary olfactory cortex neural area where olfactory info first processed, which includes the amygdale-hippocampal complex and entorhinal cortex

Amygdale-hippocampal complex conjoined regions of the amygdale and hippocampus, which are key structures in the limbic system complex is critical for unique emotional and associative properties of olfactory cognition Entorhinal cortex phylogenetically old cortical region that provides the major sensory association input into the hippocampus, the piriform cortex, and the entorhinal cortex; the limbic system is involved in many aspects of emotion and memory; olfaction is unique among the senses for its direct and intimate connection to the limbic system Limbic system encompassing group of neural structures that includes the olfactory cortex, the amygdale, the hippocampus, the piriform cortex, and the entorhinal cortex o Involved in many aspects of emotion and memory o Olfaction is unique among senses for its direct and intimate connection to the limbic system Trigeminal (V) nerves 5th pair of cranial nerves, which transmit info about the feel of an odorant (e.g. menthol feels cool, and cinnamon feels warm), as well as pain and irritation sensations (e.g. ammonia feels burning) Shape-pattern theory current dominant biochemical theory for how chemicals come to be perceived as specific odorants; shape-pattern theory contends that diff scentsas a function of odorant-shape to OR-shape fitactivate diff arrays of olfactory receptors in the olfactory epitheliathese various arrays produce specific firing patterns of neurons in the olfactory bulb, which then determine the particular scent we perceive Vibration theory alternative to shape-pattern theory, describing how olfaction works; proposes that every perceived smell has diff vibrational frequency and that molecules that produce the same vibrational freqs will smell the same Specific anosmia inability to smell one specific compound amid otherwise normal smell perception Stereoisomers isomers (molecules that can exist in diff structural forms) in which the spatial arrangement of the atoms are mirror-image rotations of one another, like R and L hand; aka optical isomers Psychophysics science of defining quantitative relationships b/w physical and psychological (subjective) events Staircase method psychophysical method for determining the concentration of a stimulus required for detection at threshold level o Stimulus (odorant) presented in ascending conc sequence until detection indicated and then conc is shifted to a descending sequence until response changes to no detection o Ascending/descending sequence is typicall repeated several times and concs at which reversals occur averaged to determine threshold detection leve of that odorant for given individual Triangle test test in which participant given 3 odors to smell, of which 2 are the samerequired to state which is odd odor out Tip-of-the-nose phenomenon inability to name an odorant, even though its very familiar; contrary to tip-of-the-tongue phenomenon, one has no lexical access to name of odorant, such

as first letter, rhyme, # of syllables, when in tip-of-the-nose state; example of how language and olfactory perception are deeply disconnected G protein-coupled receptors (GPCRs) class of receptors present on surface of OSNs; all characterized by common structural feature of 7-membrane-spanning alpha-helices Receptor adaptation biochemical phenomena that occurs after continuous exposure to an odorant, whereby the receptors stop responding to odorant and detection ceases Cross-adaptation reduction in detection of an odorant following exposure to another odorant o Presumed to occur b/c the 2 odorants share 1+ olfactory receptors for their transduction, but the order of odorant presentation also plays a role Cognitive habituation psychological process by which, after long-term exposure to an odorant, one is no longer able to detect that odorant or has very diminished detection ability Odor hedonics - linking dimension of odor perception, typically measured w/ scales pertaining to an odorants perceived pleasantness, familiarity, and intensity Gestation fetal development during pregnancy Learned taste aversion avoidance of a novel flavor after it has been paired w/ gastric illness; the smell, not the taste, of the substance is key for the learned aversion response in humans Orbitofrontal cortex part of the frontal lobe of the cortex that lies above the bone (orbit) containing the yes o Responsible for processing olfaction o Also area of brain critical for assigning affective value to sitimuli, i.e. determining hedonic meaning Main olfactory bulb (MOB) olfactory bulb; blueberry-sized extension of brain above the nose; first region of brain where smells are processed Accessory olfactory bulb (AOB) smaller neural structure located behind main olfactory bulb that receives inputs from vomeronasal organ Vomeronasal organ (VNO) chemical sensing organ at base of nasal cavity w/ curved tubular shape; VNO evolved to detect chemicals that cant be processed by olfactory epithelium, such as large and/or acqueous moleculestypes of molecules that constitute pheromones Phereomone chemical emitted by one member of a species that triggers physiological or behavioral response in another member of same species; signals for chemical communication and do not need to have any smell Releaser pheromone triggers an immediate behavioral response among conspecifics Primer pheromone triggers physiological (often hormonal) change among conspecifics; usuall involves prolonged pheromone exposure

Reading (332-357) Olfaction smell chemical detection system for molecules floating in air Gestation taste chemical detection system for molecules we put in our mouth Trigeminal system chemical sensing system important for both smell and taste experiences and innervated by trigeminal nerve

Enables us to feel gustatory and olfactory experiences, like burning and cooling

Olfactory Physiology Odors and Odorants Olfactory sensations = odors Stimuli for odors are chemical compounds, but not all chemicals are odorants to be smelled it must be volatile, small, and hydrophobic Cant smell molecules that make up the air we breathe like oxygen and nitrogen

The Human Olfactory Apparatus Olfactory system tacked onto organ that serves another purpose primary function of nose is to filter, warm, and humidify air we breath Air passes through narrow space (olfactory cleft) and settles on yellowish path of mucous membrane (olfactory epithelim retina of the nose) that contains 3 types of cells (supporting cells, basal cells, and olfactory sensory neurons, or OSNs) OSNs are small neurons with cilia protruding into mucus covering epithelium o These cilia are OSNs dendrites and have olfactory receptors (ORs) on their tips o Interaction b/w odorant and OR stimulates cascade of biochemical events, ultimately producing a.p. thats transmitted along axon of OSN olfactory bulb o Have 20 million OSNs split b/w epithelia of R and L nostrils more receptors than any other sensory system except vision o Bloodhound has 220 millions OSns and has higher proportion of brain dedicated to olfaction (5% compared to 0.1% for humans) We can smell same number of scents as dogs, but dogs can sense odors at much lower concentrations o Axons on ends of OSNs opposite the cilia (dendrites) pass through tiny sieve-like holes of cribriform plate hard blow to front of head can cause this to be fractured, slicing off fragile olfactory axons and can induce anosmia (smell blindness) = total absence of sense of smell Stem cells in olfactory epithelium can form new OSNs in fact, all OSNs die and regenerate 1x/28days but fractured cribriform plates scar over and prevent new OSN axons from passing through to brain, crippling sense of smell for life Anosmia also caused by upper respiratory infection (sinus) and sinonasal disease (polyps), medications like those treating HTN or high cholesterol, or can be congenital (3%) Recovery is best when caused by infection or disease b/c when underlying illness treated normal smell function usually returns Ppl rank loss of smell as equal to loss of big toe but anosmia can cause great suffering, and hinders taste; also profound connection b/w psychiatric depression and sense of smell ppl who lose sense of smell can fall into

clinically serious depressive states neurobiological connection b/w olfactory and emotional processing enables a bidirectional interaction b/w them 5% Americans have some olfactory dysfunction Loss can be 1st symptom of neurological discords like Alzheimers or Parkinsons In normal smell, OSN axons pass thru plate, bundle together to form olfactory nerve (Cranial Nerve I) and enter olfactory bulb where globular tangles of OSN axons synapse w/ dendrites of mitral cells and tufted cells these spherical bundles are glomeruli o All neurons expression a particular OR type, no matter where they are on nasal epithelium, convere onto 1 glomerulus pair (consisting of 1 medial and 1 lateral glomerulus) in bulb i.e. all Ors detecting menthol send their axons to single glomerulus pair thus, higher brain structures receiving info from bulbs know that signal coming from specific glomerulus pair means sniffing menthol o Olfaction is ipsilateral Central brain structures that process olfactory info from olfactory bulb are all part of network of brain structures known as limbic systems which is involved I many aspects of emotion/memory: o Primary olfactory cortex o Amygdale-hippocampal complex o Entorhibal cortex Olfactory receptor cells diff from all other receptor cells in that they arent mediated by protective barrier and make direct contract w/ brain o Many drugs can be inhaled o OSN axons thinnest and slowest in body so even w/ direct connection, time to perceive odor is longer than other senses o Lag time b/w sniff and perception ~400ms Sensation occurs when scent is neurally registered, perception occurs when we become aware of detecting the scent

Genetic Basis of Olfactory Receptors In general, the more receptors that are expressed from genome (we only express about 3040%), the more sensitive one is to odorants Genetic variation in receptor expression may be the one innate factor in the modulation of our odor likes and dislikes High proportion of OR pseudogenes in humans could be result of evolutionary trade-off b/w vision and olfaction payoff from superior visual detection must have been better for our ancestors survival than disadvantages from diminished olfactory acuity

The Feel of Scent Most odorants stimulate the somatosensory system to some degree through polymodal nociceptors (touch, pain, and temp receptors) inside the nose o E.g. menthol = cool, ammonia = burning

These sensations mediated by trigeminal nerve (Cranial Nerve V) Nasal cooling (CN V) and specific scent (CN I) associated w/ smell of peppermint fuse to produce holistic sensory experience Why we cry when chopping onions or sneeze sniffing pepper

From Chemicals to Smells How does biochemical interaction b/w an odorant and an OR, and subsequent neurological processing in the olfactory bulbs and later brain structures, result in the psychological perception of a scent like a lemon?

Theories of Olfactory Perception Shape-pattern theory contends that odorant molecules have diff shapes and Ors diff shapes and odorant detected by specific OR to the extent that the odorants molecules fit into OR when odorant snifferd, particular pattern generated across glomeruli and differences in those spatial patterns provide basis for array of odors we perceive o Combinatorial code diff scents activate diff arrays of ORs in epithelia producing specific firing pattern of neurons in bulb and this pattern of electrical activity then determines scent perceived Alternative theory = vibration theory b/c of atomic structure, each molecule has diff vibrational freq and molecules producing same freqs have same smell various chemicals w/ similar vibrations due to molecule compositions have similar smells o This cant explain specific anosmias and different scents produced by stereoisomers which shape-pattern theory can explain Specific anosmia = inability to smell 1 specific compound amid otherwise normal smell perception o Likely genetic o Inability to smell androstenone in sweat and pork 50% of pop have this specific anosmia o Study shows variability in detection of androstenone as well as its perceived pleasantness is due to genetic diffs in OR expression b/w individuals o Able to induce sensitivity to androstenone through repeated exposure Plasticity and modulation through environmental influences appear to be basic principles in olfaction

The Importance of Patterns As with detecting colors, we detect odors by considering the pattern of activity across various diff receptor types Receptor activation also affected by odor intensity, which helps explain why dogs w/ many more functional receptors than we have can perceive considerably lower concs of odorants Also order of OR activation important thus diff odor perceptions can be due to diff OR firing patterns or to firing of same receptors at diff rate or sequence

Odor Mixtures Rarely smell pure odorants outside of labs How do we process components in an odorant mixture? o Analysis audition uses this when high and low note played and we perceive them as separate o Synthesis colors use this when we mix red and green lights and see yellow THIS IS FOR ODORS most mixtures perceived as unitary wholes o Olfaction primarily synthetic sense but a certain amount of analytical ability can be developed w/ training

Odor Imagery Degree of overlap between smelling an odor and imaging it (trying to reproduce smell of odor in minds nose) is weak Dreams w/ olfactory sensations also rare b/cwe dont think in smell, its not necessary to have stored representations of olfactory experiences but, since olfactory system appears to be highly flexible and modulated by learning and enviro experiences throughout life, you may be able to develop capacity to create sensory representations of smells w/ training

Olfactory Psychophysics, Identification, and Adaptation Goal of psychophysicist is to quantify the psychological experience of our sensory orld

Detection, Discrimination, and Recognition How much stimulation is required before we perceive something to be there? o Olfactory detection thresholds depend on number of factors Longer carbon chains easier to detect Women have lower thresholds than men, especially during ovulatory period of menstrual cycles (though not heightened during pregnancy) Detection decreases w/ age b/c change in proportion of OR cell regeneration to cell death as we age, # ORs that die off continues to rise beyond # regenerated > age 85, ~50% population is anosmic, though taste and trigeminal perception dont decline w/ age Can discriminate thousands of odors and the more training the more we can discriminate Recognition is ability to remember whether or not weve smelled an odor before o Takes 3x as many odorant molecules to go through nose to regoxnize an odor than to simply detect that odor is there

Durability of odor 30-second delay b/w odor presentation and testing produces big drop in recognition accuracy but what we remember after 30s pretty close to what we remember after 3 days, month, or even a year o Memory of odors especially potent if emotion is experienced during initial exposure

Psychophysical Methods for Detection and Discrimination Staircase method used to determine odor detection threshold (definition more in-depth) Triangle test used to determine if someone can discriminate b/w 2 odors (def has in-depth)

Identification Called mute sense since we are often lost for verbal descriptors for olfactory experiences Tip-of-the-nose phenomenon is diff than of tongue b/c we typically know nothing about the label were searching for but even if we cant name it we often know how to respond to it, i.e. we smell maple syrup and wed like to eat it Fewer words that refer exclusively to our experience of smells than for any other sensation Unlike other senses, olfactory info not integrated in thalamus prior to cortex processing and argued that thalamus has relevance for language Majority of olfactory processing in R hemi, but language in L hemi New studies show odors and language processing may compete for cognitive resources that share the same neural substrates Recognition tests like scratch and sniff important for detecting early stages of neurological disorders like Alzheimers b/c

Adaptation Sense of smell is change detector system When odorant molecules bind to corresponding OSNs in nose, the ORs retreat into cell body and receptors are no longer physically available to respond to same odorant o Known as receptor recyclying o When OR internalized by cell body, the odorant becomes unbound and is then recycled through cell and emerges again in number of minutes o Common to all receptors in class of G protein-coupled receptors (GPCRs) Process = receptor adaptation On avg, 15-20 mins of continuous exposure to an odorant for molecules to stop eliciting an olfactory response, but adaptation can also occur in <1 min Adaptation can be undone quickly by stepping away from scent Magnitude of adaptation affected by odor intensityas odor conc increases, degree of adaptation decreases Psychological processes also can have effect on adaptation rates o When told odorant healthful adaptation took place within 20 mins o When told odorant hazardous ppl more sensitized than at start of experiment Benefit = filter out stable background odors

Active sniffing makes OR neurons less responsive to stable odors and more responsive to new ones Sometimes, exposure to one odorant can raise detection threshold for second odorant = crossadaptation o Presumed to occur when odorants rely on similar sets of ORs o However, complicated by act that cross-adaptation relationships are nonreciprocal o Go away after a few minutes

Cognitive Habituation Cognitive habituation is psychological effect of no longer reacting or responding to odor we live with o Requires wks to reversewont smell cologne again after 5 daysneed to abstain >2wks 3 possible mechanisms involved, either singularly or in interaction o 1) ORs that are internalized in cell bodies during adaptation may be more hindered after continuous exposure and take much longer to recycle than usual o 2) continuous exposure could lead to odorant molecules absorbing into bloodstream, then transported to ORs via nasal capillaries when we breathe out through noseas long as in bloodstream, we will be constantly adapted o 3) cognitive-emotional factors can be involved in reverse direction (told something harmful and participants kept smelling but reverse of this) Cant smell when asleep do not awaken or have EEG sleep pattern changes even w/ potent odorants fMRI study shows odors may be detected by brain during SWS (slow-wave sleep) could be that since attention is cut off during sleep so is our ability to respond to odors so although odors may be registered by brain without conscious awareness (sensation), olfactory perception depends on attention

Olfactory Hedonics Odor hedonics = affective evaluations of odors, i.e. like or dislike

Familiarity and Intensity We like odors weve smelled before Perceive pleasant odors as familiar Intensity related to odor liking often has inverted-U functionscent rated as more positive with increasing intensity to a point at which function reverses and stronger it is, the less pleasant Other times a bad smell just gets worse with intensity linear downward

Nature or Nurture? Innate/nature o We are born w/ predisposition to like/dislike various smells

Researchers found that molecular structures of odorants was able to account for 30% of variance in participants responses so odor pleasantness perception may be innate Learned/nurture o Born merely w/ predisposition to like/dislike smells and that whether smell is liked or not is determined by the emotional value (good or bad) of experiences that have been associated with it o Cultural learning also provides meaning to many unencountered stimuli like skunk o LOTS OF EVIDENCE HERE o Infants found that when presented w/ odorants they havent encountered before, infants/children often display very diff preferences from those of adults dont find sweat or feces unpleasant and toddlers dont hedonically differentiate b/w odorants adults find very unpleasant or pleasant o Study difficulty olfactory system fully functional by 3rd month of gestation and odorant molecules find way into womb o Fetus studies mothers eating distinctive-smelling volatiles (garlic, cigarettes) during pregnancy or breast-feeding had infants w/ greater preferences for these smells than those who hadn t been exposed o Cross-cultural data culturally-polarized responses to specific odors Asians hate smell of cheese and we hate fermented soybean dish even though they are similar in protein and nutrition content Some African cultures like smell of feces Impossible to find odor for stink bomb that was unanimously considered repellent across ethnic grps o Novel odorant experiments can be made to be perceived as good or bad as a function of the associations made to it

An Evolutionary Argument Supports learned-odor preference theory Specialist animal species live in specific habitats and so have limited # food sources and predators, thus hardwired responses to particular odors are adaptive Generalist species (like humans) can exploit many diff habitats and resources/predators vary widely across enviros so not adaptive for these species to have predetermined olfactory responses to any particular odor thus need to be ready to learn and remember what to approach/avoid based on experience o Evidence that learning is critical mechanism for generalists to acquire odor responses shown by learned taste aversions Made to avoid flavor by being made sick after rats w/ drink and children w/ new ice cream pre-chemo

Caveats 1) Trigeminally irritating odors may elicit pain responses and we all have innate drive to avoid pain 2) Potential variability in receptor genes and pseudogenes that are expressed across individuals may influence odor intensity perception and consequently pleasantness, i.e. perceived intensity of scent may be partly responsible for its perceived (un)pleasantness o Genetic differences in OR expression appear to extend to ethnicities as well which may help explain why its been impossible to develop a universally effective stink bomb

Olfaction, Memory, and Emotion In British study, wintergreen given one of lowest pleasantness ratings but in American, rated the most pleasant there is historic reason behind this in Britain, wintergreen associated w/ medicine whereas in US it is exclusively a candy smell so only has sweet, positive connotations o The key to olfactory associative learning is the experience that occurs when odor is first encountered and in particular, the emotional connotations of that experience Are odors best cues to memory? o STUDY compared recollections stimulated by popcorn w/ memories evoked by sight or it, sound of popping, feel of kernels, or word popcorn In terms of vividness and accuracy, memories evoked by odors are good, but not any better than-memories evoked by sights, sounds, feel, or words o BUT, odor cues are distinctive in their emotionality STUDIES participants list more emotions w/ olfactory cues than they do w/ other sensory modalities, rate emotions as having greater intensity, and report particular memories as being more emotionally laden when elicited by odors; also feel transported to original time and place when scent elicits recall than when memory triggered via another sensory system Odors no better than other cues at eliciting accurate info, but emotion and evocativeness of odor-elicited memories leads to false impression that such memories are especially accurate

Neuroanatomical and Evolutionary Connections between Odor and Emotion Amygdale which synapses directly w/ olfactory nerve is critical for emotional associative learning o STUDY more activated when recalling memory connected w/ smell of perfume vs. sight of it or when smelled or saw nonmeaningful perfume Orbitofrontal cortex (where olfaction is processed) is also cortical area responsible for assigning affective value, or hedonic judgments in response to wide range of stimuli Rhinencephalon (nose-brain) comprises the olfactory cortex and limbic areas developed first from olfactory structures and only later in evolution did limbic structures like amygdale develop maybe our hedonic and emotional rxns to stimuli in general have origin in sense of smell

Emotions and olfaction are functionally analogousapproach or avoidboth enable organism to react appropriately to its enviro, maximizing chances for basic survival and reproductive success o In this way, human emotional system can be seen as highly evolved, abstract cognitive version of basic behavioral motivations instigated by olfactory system in animals

The Vomeronasal Organ and the Question of Human Pheromones For animals that rely on smell to survive, olfactory system has 2 subdivisions o Main olfactory bulb (MOB) o Accessory olfactory bulb (AOB) add on to back of MOB o Neurons from these 2 systems dont interconnect and function separately o To activate AOB, structure diff from nose needs to be engaged, known as vomeronasal organ (VNO) VNO can respond to some olfactory stimuli but primarily responds to chemicals that are higher in molecular weight than can be detected by OSNs as well as to nonvolatile chemicals and those that dissolve in water o Humans dont have a VNO or AOB o Primary function o detect pheromones, which arent odors but are chemicals that may or may not have a smell Pheromones = chemical communication chemical compound produced by one animal that elicits a specific behavioral or physiological response in another animals of same species o Mark territory, alarm or defense rxns, mating behavior o Produce 2 kinds of effects Releaser pheromones fast, always produce behavioral responses Primer pheromones slow, produce physiological change in recipient over time In humans, there is McClintock effect- women in physical proximity over time start to have coinciding menstrual cycles o Believed to be caused by chemicals present in human sweat capable of priming hormonal systems of other individuals o Female body odors that vary w/ hormonal status appear to influence human sexual arousal