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quantified by measuring intra-nasal pressure or flow or air or, while less accurate, through a strain gauge on the chest to measure total respiratory volume. Strategies for sniffing behavior vary depending upon the animal, with small animals (rats, mice, hamsters) displaying sniffing frequencies ranging from 4 to 12 Hz but larger animals (humans) sniffing at much lower frequencies, usually less than 2 Hz. Subserving sniffing behaviors, evidence for an "olfactomotor" circuit in the brain exists, wherein perception or expectation of an odor can trigger brain
75:
276:, or a pressure sensor. These can be inserted temporarily into the nares or implanted surgically. The basic principles of operation are shared between the temperature and pressure devices. Inhalation of ambient air provides cool temperature into the nasal cavity, whereas exhalation of inhaled air provides warm temperature into the nasal cavity and simultaneously an increase in intranasal pressure as air from the lungs is forced out of the nostrils. Placement of these sensors close to the
27:
396:, in the ability for both methods to provide rapid "snapshots" of information to the brain. This analogy, though, may be imprecise since small animals (e.g., mice) make odor-based decisions (through sniffing) while also making visual decisions, yet do not saccade. Sniffing is also fundamentally similar to active touch, including swiping ones finger along a surface to scan texture.
78:
82:
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264:. This method has advantages to directly index air leaving the nares (increase in microphone output), yet is mostly non-invasive. Due to this non-invasive nature of microphone measures, these methods have been employed in dogs during odor tracking exercises and are useful for measuring sniffing on a temporary basis in other large animals.
80:
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brain is temporally linked to the respiratory cycle, with bouts of activity occurring with each inhalation. This linkage between sniffing frequency and odor processing provides a mechanism for the control of odor input into the brain by respiratory frequency and possibly amplitude, though this is not well established.
325:) also display sniffing behaviors during underwater odor-guided tasks. Shrews inhale-exhale small amounts of air in a precise and coordinated fashion while tracking an underwater odor trail. This occurs through the inhalation of air above ground, to allow air to volatilize odors in an environment otherwise void of air.
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sniffing. This was the first paper to report that rats can sniff at frequencies reaching 12 Hz upon detection of odors and during free exploration. This paper also provided early evidence that the rhythm of sniffing was coupled with other sensory behaviors, such as whisking, or the movement of the whiskers.
403:
responsible for governing some other active sampling behaviors, sniffing in animals often occurs at similar frequencies (2 to 12 Hz) and in a phasic relationship to the active sampling behaviors of whisking and licking. Whisking and sniffing are tightly correlated in their occurrence, with sniff
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Transitions in sniffing frequency are observed in animals performing odor-guided tasks. Studies of recording sniffing in the context of odor-guided tasks involve implanting intranasal temperature and pressure sensors into the nasal cavity of animals and either measuring odor-orienting responses (fast
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Perhaps the simplest method for determining the moment of sniffing is video-based. High resolution video of small animals (e.g., rats) during immobile respiration enables approximations of sniffing, including identification of individual sniff events. Similar methods can be employed to identify fast,
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One of the earliest reports of exploring sniffing in non-human animals was provided by Welker in his 1964 article, Analysis of sniffing in the albino rat. In this study, Welker used video recordings of rats during presentation with odors and other stimuli to explore the chest movements as an index of
300:
using video-based measures. In this study robust changes in respiratory frequency were reported to occur during exploration of an open arena and novel odors. Resting respiration occurs ~2 times/second (Hz), and increases to about 12 Hz are noted during states of exploration and arousal. Similar
188:
within the hedgehog olfactory bulb were entrained to the respiratory cycle. Further, odor-evoked oscillations (including an exhaled puff from a pipe), were amplified along with the respiratory cycle. These data gave evidence that information processing within the brain, particularly that of odors,
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While sniffing behavior is often observed and discussed within the context of acquiring odor information, sniffing is also displayed during the performance of motivated behaviors and upon deep brain electrical stimulation of brain reward centers. For instance, prior to obtaining a food reward, mice
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Notably, several studies have reported that modulation in sniffing frequency may be just as great in context of anticipation of odor sampling as during sampling of odors. Similar changes in sniffing frequency are even seen in animals presented with novel auditory stimuli, suggesting a relationship
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and the frequency of inhalations. Both of these entail modulations in the manner whereby air flows within the nasal cavity and through the nostrils. As a consequence, when the air being breathed is odorized, odors can enter and leave the nasal cavity with each sniff. The same applies regardless of
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in humans and in fact, in humans, a single sniff is often sufficient for optimal odor perception. For instance, a deep, steady inhalation of a faint odor allows a more potent percept than a shallow inhalation. Similarly, more frequent sniffs provide a faster percept of the odor environment than
163:
Studies into the perceptual correlates of sniffing on human olfaction did not reach the mainstream scientific community until the 1950s. Frank Jones, an
American psychologist, published a paper demonstrating the interplay between parameters of sniffing and odor detection thresholds. He found that
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studies lack resolution to determine the impacts of sniffing frequency on the structure of odor input through the brain, although imaging studies have revealed that the motor act of sniffing is anatomically independent of sniff-evoked odor perception. Implications for this include the shared but
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Measurements of sniffing simultaneously with physiological measures from olfactory centers in the brain have provided information on how sniffing modulates the access and processing of odors at the neural level. Inhalation is necessary for odor input to the brain. Further, odor input through the
201:
in the nose and consequentially odor input to the brain. Later, evidence that single neurons in the olfactory bulb, the brain's first relay station for odor information, are entrained with respiration was presented, establishing a solid basis for the control of odor input to the brain and the
44:
for the purpose of information acquisition. This behavior, displayed by all terrestrial vertebrates, is typically identified based upon changes in respiratory frequency and/or amplitude, and is often studied in the context of odor guided behaviors and olfactory perceptual tasks. Sniffing is
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The rapid modulation of sniffing upon inhalation of a novel odor or an irritating odor is evidence for an "olfactomotor" loop in the brain. In this loop, novel odor-evoked sniffing behavior can occur rapidly upon perception of a novel odor, one of interest, or an odor which is aversive.
171:
While behavioral and psycho-physical studies into sniffing and its influence on odor perception began to surface, much less work was being performed to explore the influence of sniffing behaviors on the physiological processing of odors within the brain. Early recordings from the
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and rabbits increase their sniffing frequency in a manner independent of seeking odor information. Sniffing behavior is also displayed by animals upon involuntary electrical stimulation of numerous brain structures. Thus, while sniffing is often considered a critical part of
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inhalations occurring during whisker protraction. Due to the metabolic need to coordinate breathing and swallowing, small animals (rats and mice) often lick at similar frequencies of sniffing (4 to 8 Hz) and swallow in between inhalations or during brief periods of
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on several grounds. In humans, one can assess the occurrence of a sniff based upon volitional control of air movement through the nose. In these cases, human subjects can be asked to inhale for a certain amount of time, or in a particular pattern. Some animals are
375:
which governs inhalation/exhalation patterns. Activity from respiratory brain stem structures then modulates nervous activity to control lung contraction. To exert changes to respiration, and thereby evoke sniffing behavior, volitional centers in the
210:
There are multiple methods available for measuring sniffing. While these methods are applicable for most animal models (mice to humans), selection of appropriate sniff measurement methods should be determined by experimental need for precision.
132:, wherein the only air for respiration must arrive into the lungs via the nose. This includes rats and mice. Thus, in these animals the distinction between a breath and a sniff is not clear and could be argued to be indistinguishable. (See
347:
only sniffing once every 3 seconds. These examples have been supported by empirical studies (see above) and have provided insights into methods whereby humans may change their sniffing strategies to modulate odor perception.
79:
65:
movements in small animals (viz., whisking). Atypical sniffing has been reported in cases of neurological disorders, especially those disorders characterized by impaired motor function and olfactory perception.
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Sensors to measure chest expansion during inhalation provide direct information of sniff cycles. These methods include mechanical and optical devices. Mechanical devices for sniffing measurements are
139:
Sniffing is observed among all terrestrial vertebrates, wherein they inhale environmental air. Sniffing may also occur in underwater environments wherein an animal may exhale air from within its
392:
Sniffing, as an active sampling behavior, is often grouped along with other behaviors utilized to acquire sensory stimuli. For instance, sniffing has been compared to rapid eye movements, or
301:
transitions in sniffing frequency are observed in freely exploring mice, which, however, maintain generally higher sniffing frequencies than rats (3 to 15 Hz vs 2 to 12 Hz).
424:
have abnormal sniffing capabilities (i.e., reduced volume and flow rate) which may underlie olfactory perceptual impairments in the disease. Studies into sniffing in mouse models of
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into an air-tight chamber with a pressure transducer embedded within to access nasal transients, while simultaneously odors are presented to measure responses while nose-poking.
240:
around the chests of larger animals. In both cases, a positive increase in signal output (voltage) can be identified and used to index inhalation events. Alternatively, a photo
561:
Youngentob, S.L.; Mozell, M. M.; Sheehe, P. R.; Hornung, D. E. (1987). "A quantitative analysis of sniffing strategies in rats performing odor discrimination tasks".
671:
Verhagen, J. V.; Wesson, D. W.; Netoff, T. I.; White, J. A.; Wachowiak, M (2007). "Sniffing controls an adaptive filter of sensory input to the olfactory bulb".
224:
and stimulus investigation. This method, however, does not provide direct evidence for sniffing and is not reliable in larger animals (rabbits to humans).
248:). In this design, a decrease in signal reflects inhalation (chest expansion) as the chest would interrupt the light passage to the photo transducer.
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was linked with respiration - establishing the integral nature of sniffing for the physiological processing of odors. About 20 years later,
1725:
Gottfried, J.A.; Winston, J. S.; Dolan, R. J. (2006). "Dissociable codes of odor quality and odorant structure in human piriform cortex".
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Massengale, O.N.; Glaser, H. H.; LeLievre, R. E.; Dodds, J. B.; Klogk, M. E. (1963). "Physical and
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Johnson, B. N.; Mainland, J. D.; Sobel, N. (2003). "Rapid olfactory processing implicates subcortical control of an olfactomotor system".
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Donald A. Wilson and
Richard J. Stevenson Learning to Smell: Olfactory Perception from Neurobiology to Behavior Johns Hopkins Press, 2006
272:
The most precise methods to date to measure sniffing involve direct intranasal measures through use of a temperature probe, called a
2288:
Gordon M. Shepherd
Neurogastronomy: How the Brain Creates Flavor and Why It Matters New York : Columbia University Press, 2012
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Uchida, N.; Kepecs, A.; Mainen, Z. F. (2006). "Seeing at a glance, smelling in a whiff: rapid forms of perceptual decision making".
305:
sniffing) or sniffing during performance in operant odor-guided tasks. Alternatively, animals can be conditioned to insert their
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1927:
Kepecs, A.; Uchida, N.; Mainen, Z. F. (2007). "Rapid and precise control of sniffing during olfactory discrimination in rats".
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Sobel, N; Thomason, M.E; Stappen, I.; Tanner, C. M.; Tetrud, J. W.; Bower, J. M.; Sullivan, E. V.; Gabrieli, J. D. E. (2001).
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Vanderwolf, C. H. (2001). "The hippocampus as an olfacto-motor mechanism: were the classical anatomists right after all?".
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and also humans have not found major effects of
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Macrides, F.; Chorover, S. L (1972). "Olfactory bulb units: activity correlated with inhalation cycles and odor quality".
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380:
must stimulate brain stem structures. It is through this simple pathway that the decision to inhale or sniff may occur.
57:
and thus acquisition of odor information. Sniffing is analogous to other stimulus sampling behaviors, including visual
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Vanderwolf, C. H. (1992). "Hippocampal activity, olfaction, and sniffing: an olfactory input to the dentate gyrus".
768:"Temporal relationship between sniffing and the limbic theta rhythm during odor discrimination reversal learning"
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As a direct measurement of sniffing, early studies favored the use of microphones placed/secured external to the
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2520:
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Laing, D.G. (1986). "Identification of single dissimilar odors is achieved by humans with a single sniff".
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1936:
425:
421:
164:
deep sniffs, consisting of a large volume of air, allowed for consistent and accurate detection of odors.
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While sniffing is generally thought to occur solely in terrestrial animals, semi-aquatic rodents (
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and nasal cavity to acquire odors within an aquatic environment and then re-inhale this air. (See
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on sniffing behavior, although numerous neurological disorders affect respiration. Humans with
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155:, its link with motivated and reward behaviors suggests it plays a role in other behaviors.
2069:"Pre-Bötzinger complex: a brainstem region that may generate respiratory rhythm in mammals"
1163:"An impairment in sniffing contributes to the olfactory impairment in Parkinson's disease"
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26:
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is a perceptually-relevant behavior, defined as the active sampling of odors through the
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In part due to the interrelatedness of the respiratory brain stem structures with other
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and solvents, and other industrial chemicals which may be inhaled as a form of drug or
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Smith, J.C.; Ellenberger, H. H.; Ballanyi, K.; Richter, D. W.; Feldman, J. L. (1991).
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can be placed on the opposite side of an animal's chest from a light source (e.g., a
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published a series of studies wherein he further proposed that the flow rate and the
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2194:
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1006:
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The earliest published study of sniffing behavior in small animals was performed in
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properties of odorants interplay to affect the location of odorant binding to
108:
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1989:
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1309:"Perceived effort in sniffing: The effects of sniff pressure and resistance"
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50:
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1225:"Respiratory and sniffing behaviors throughout adulthood and aging in mice"
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Odor inhalation evokes activity throughout olfactory structures in humans.
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2007:
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58:
1877:"Associative Cortex Features in the First Olfactory Brain Relay Station"
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of animals allows measures of odorized air transients as they reach the
2468:"Interplay between sniffing and odorant sorptive properties in the rat"
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62:
20:
2306:
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1974:"The control of responding by sounds: unusual effect of reinforcement"
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1443:
497:
284:
and thus are common methods for measuring sniffing in the context of
113:
103:
The behavior of sniffing incorporates changes in air flow within the
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716:"Speed and accuracy of olfactory discrimination in the rat"
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for their work on the functions of neurons, revealed that
2545:
2425:"All in a sniff: olfaction as a model for active sensing"
1826:"Rapid encoding and perception of novel odors in the rat"
388:
Relation of sniffing to other stimulus sampling behaviors
1475:
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distributed pathways for odor processing in the brain.
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180:, who previously won the 1932 Nobel Prize along with
236:foils placed under the chests of small animals and
1573:"Olfactory reactions in the brain of the hedgehog"
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2556:Science Daily: Olfaction and Alzheimer's disease
2208:Li, W.; Howard, J. D.; Gottfried, J. A. (2010).
1778:Thesen, A.; Steen, J. B.; Doving, K. B. (1993).
1109:Deschenes, M.; Moore, J.; Kleinfeld, D. (2011).
123:The act of sniffing is considered distinct from
329:Sniffing and control of odor input to the brain
2378:Cheung M.C.; Carey R.M.; Wachowiak M. (2009).
972:
970:
144:
133:
1780:"Behaviour of dogs during olfactory tracking"
1669:
1667:
8:
2308:"The world smells different to each nostril"
2126:"The sniff is part of the olfactory percept"
1417:
1415:
556:
554:
552:
268:Nasal thermocouple and nasal pressure sensor
2531:Case Western Reserve University, Wesson Lab
1922:
1920:
1819:
1817:
1815:
1346:
1344:
107:. This can involve changes in the depth of
2019:
2017:
1523:
1521:
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1307:Teghtsoonian, R.; Teghtsoonian, M (1982).
1218:
1216:
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915:Sobel, N.; V. Prabhakaran; J. E. Desmond;
761:
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220:high frequency sniffing during states of
19:For the act of clearing nasal mucus, see
524:
522:
520:
518:
516:
514:
512:
416:Few studies have explored the impact of
363:Sniffing is fundamentally controlled by
49:to allow for the modulation of sniffing
2516:Association for Chemoreception Sciences
910:
908:
906:
819:
817:
815:
813:
811:
508:
342:The nature of sniffing regulates odor
112:what gas is being inhaled, including
7:
2351:The Journal of Experimental Biology
1477:. Academic Press. pp. 273–282.
412:Relevance to neurological disorders
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1875:Doucette, W.; et al. (2011).
1111:"Sniffing and whisking in rodents"
784:10.1523/JNEUROSCI.02-12-01705.1982
70:Background and history of sniffing
14:
202:processing of odors by sniffing.
2124:Mainland, J.; Sobel, N. (2006).
714:Uchida, N; Mainen, Z. F (2003).
317:Sniffing in semi-aquatic animals
206:Methods for quantifying sniffing
16:Nasal inhalation to sample odors
1784:Journal of Experimental Biology
1274:New England Journal of Medicine
1115:Current Opinion in Neurobiology
919:; R. L. Goode; E. V. Sullivan;
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2551:Olfactory-auditroy integration
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1589:10.1113/jphysiol.1942.sp003955
1313:Perception & Psychophysics
313:between sniffing and arousal.
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2472:The Journal of Neuroscience
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288:and psychological studies.
2597:
1229:Behavioural Brain Research
1127:10.1016/j.conb.2011.11.013
408:(cessation of breathing).
401:central pattern generators
359:Neural control of sniffing
199:olfactory receptor neurons
18:
2526:The LRI and Odor Database
1748:21.11116/0000-0001-A18C-B
1241:10.1016/j.bbr.2011.04.016
292:Sniffing in small animals
145:sniffing in small animals
134:sniffing in small animals
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130:obligate nasal breathers
91:sniffing the ground and
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1972:Harrison, J.M. (1979).
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2581:Respiratory physiology
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2143:10.1093/chemse/bjj012
1951:10.1152/jn.00071.2007
1797:10.1242/jeb.180.1.247
1577:Journal of Physiology
1571:Adrian, E.D. (1942).
1473:Karpov, A.P. (1980).
1369:10.1152/jn.90981.2008
1034:10.1152/jn.00115.2003
631:10.1093/chemse/bjn029
373:Pre-Botzinger complex
176:of hedgehogs by Lord
86:
29:
2423:Wachowiak M (2011).
2357:(Pt 24): 3891–3902.
2227:10.1093/brain/awq209
463:Nasal administration
323:American water shrew
286:sensory neuroscience
278:olfactory epithelium
246:Light-emitting diode
61:, active touch, and
2478:(44): 15577–15589.
2085:1991Sci...254..726S
1688:1972Sci...175...84M
1634:1964Natur.203.1181M
1628:(4950): 1181–1182.
1436:2006Natur.444.1024C
1430:(7122): 1024–1025.
1179:2001PNAS...98.4154S
937:1998Natur.392..282S
426:Alzheimer's disease
422:Parkinson's disease
365:respiratory centers
282:olfactory receptors
186:neural oscillations
32:sniffing each other
1326:10.3758/bf03202655
483:Olfactory tubercle
453:History of perfume
338:Sniffing in humans
96:
47:respiratory center
35:
2294:978-0-231-15910-4
2284:978-1-4200-7197-9
2079:(5032): 726–729.
1642:10.1038/2031181a0
1280:(25): 1340–1344.
931:(6673): 282–286.
921:J. D. E. Gabrieli
778:(12): 1705–1711.
726:(11): 1224–1229.
458:Machine olfaction
84:
2588:
2505:
2495:
2462:
2452:
2419:
2409:
2399:
2374:
2345:
2327:
2250:
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2220:(9): 2714–2726.
2205:
2199:
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2156:
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2145:
2121:
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2114:
2104:
2064:
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2012:
2011:
2001:
1978:J Exp Anal Behav
1969:
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1944:
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1887:(6): 1176–1187.
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1363:(2): 1089–1102.
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1328:
1304:
1298:
1297:
1269:
1263:
1262:
1252:
1220:
1211:
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1200:
1190:
1173:(7): 4154–4159.
1158:
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1138:
1106:
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1099:
1068:Nat Rev Neurosci
1063:
1054:
1053:
1028:(2): 1084–1094.
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610:
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537:(3–4): 223–244.
526:
478:Olfactory system
371:, including the
252:Nasal microphone
85:
2596:
2595:
2591:
2590:
2589:
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2384:Chemical Senses
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2256:Further reading
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493:Piriform cortex
448:Electronic nose
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378:cerebral cortex
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298:laboratory rats
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2013:
1984:(2): 167–181.
1964:
1935:(1): 205–213.
1929:J Neurophysiol
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1022:J Neurophysiol
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869:Brain Research
859:
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733:10.1038/nn1142
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685:10.1038/nn1892
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1235:(1): 99–106.
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28:
22:
2475:
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2390:(1): 63–76.
2387:
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2354:
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2318:(6757): 35.
2315:
2311:
2300:Publications
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775:
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723:
720:Nat Neurosci
719:
709:
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673:Nat Neurosci
672:
622:
618:
569:(1): 59–69.
566:
562:
534:
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473:Olfactometer
415:
398:
391:
382:
362:
352:Neuroimaging
349:
341:
332:
320:
311:
303:
295:
274:thermocouple
271:
262:nasal cavity
255:
238:strain gauge
231:
228:Chest strain
218:
209:
178:Edgar Adrian
170:
166:
162:
149:
138:
122:
102:
42:nasal cavity
37:
36:
2576:Respiration
2130:Chem Senses
1790:: 247–251.
619:Chem Senses
125:respiration
2570:Categories
2561:Odor Space
1836:(4): e82.
1408:: 159–176.
1121:(2): 1–8.
826:Perception
772:J Neurosci
504:References
488:Phantosmia
438:Inhalation
344:perception
242:transducer
191:Max Mozell
109:inhalation
99:Background
89:island fox
1937:CiteSeerX
1830:PLOS Biol
961:205002040
750:205430292
531:Behaviour
369:brainstem
153:olfaction
55:amplitude
51:frequency
30:Two dogs
2502:23115193
2459:21943596
2416:18791186
2371:12432011
2334:10573415
2246:20724290
2195:19717010
2152:16339268
2054:44901866
1959:17460109
1911:21435561
1862:18399719
1765:13906088
1757:16446149
1712:33998763
1650:14213677
1607:16991539
1558:14376691
1452:17183311
1387:19052108
1294:14064317
1259:21524667
1207:11259673
1145:22177596
1088:16715056
1050:16446791
1042:12711718
1007:21832964
999:11718883
897:12799792
854:31619356
742:14566341
701:15324047
693:17450136
649:18534995
591:43798119
432:See also
394:saccades
195:sorption
59:saccades
38:Sniffing
2493:3495330
2450:3237116
2407:2639450
2342:4416272
2237:2948816
2187:9809310
2111:1683005
2102:3209964
2081:Bibcode
2073:Science
2046:3737714
1999:1332893
1902:3064824
1853:2288628
1806:8371085
1704:5008584
1684:Bibcode
1676:Science
1658:4208991
1630:Bibcode
1598:1393326
1550:1418901
1510:5164362
1460:4417227
1432:Bibcode
1378:2657070
1335:7110885
1250:3128824
1175:Bibcode
1136:4934665
1096:5893980
953:9521322
933:Bibcode
889:1450928
846:6657430
802:7143047
793:6564372
640:2533419
583:3685154
367:in the
222:arousal
159:History
63:whisker
21:Sniffle
2500:
2490:
2457:
2447:
2429:Neuron
2414:
2404:
2369:
2340:
2332:
2312:Nature
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2193:
2185:
2150:
2109:
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2008:501270
2006:
1996:
1957:
1939:
1909:
1899:
1881:Neuron
1860:
1850:
1804:
1763:
1755:
1727:Neuron
1710:
1702:
1656:
1648:
1622:Nature
1605:
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1424:Nature
1385:
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925:Nature
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748:
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699:
691:
647:
637:
589:
581:
498:Smound
307:snouts
114:toxins
2338:S2CID
2261:Books
2214:Brain
2191:S2CID
2050:S2CID
1761:S2CID
1708:S2CID
1654:S2CID
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