Nasal inhalation does not improve memory of visual repetitions.

Several studies suggest that breathing entrains neural oscillations and thereby improves visual detection and memory performance during nasal inhalation. However, the evidence for this association is mixed, with some studies finding no, minor, or opposite effects. Here, we tested whether nasal breathing phase influences memory of repeated images presented in a rapid serial visual presentation (RSVP) task. The RSVP task is ideal for studying the effects of respiratory-entrained oscillations on visual memory because it engages critical aspects of sensory encoding that depend on oscillatory activity, such as fast processing of natural images, repetition detection, memory encoding, and retrieval. It also enables the presentation of a large number of stimuli during each phase of the breathing cycle. In two separate experiments (n = 72 and n = 142, respectively) where participants were explicitly asked to breathe through their nose, we found that nasal breathing phase at target presentation did not significantly affect memory performance. An exploratory analysis in the first experiment suggested a potential benefit for targets appearing approximately 1 s after inhalation. However, this finding was not replicated in the pre-registered second experiment with a larger sample. Thus, in two large sample experiments, we found no measurable impact of breathing phase on memory performance in the RSVP task. These results suggest that the natural breathing cycle does not have a significant impact on memory for repeated images and raise doubts about the idea that visual memory is broadly affected by the breathing phase.

Metabolic state modulates neural processing of odors in the human olfactory bulb.

The olfactory and endocrine systems have recently been shown to reciprocally shape the homeostatic processes of energy intake. As demonstrated in animal models, the individual's metabolic state dynamically modulates how the olfactory bulb process odor stimuli using a range of endocrine signals. Here we aimed to determine whether the neural processing of odors in human olfactory bulb is modulated by metabolic state. Participants were exposed to food-associated odors, in separate sessions being hungry and sated, while neural responses from the olfactory bulb was obtained using electrobulbogram. We found significantly higher gamma power activity (51-100 Hz) in the OB's response to odors during the Hunger compared to Sated condition. Specifically, EBG gamma power were elevated while hungry already at 100 ms after odor onset, thereby suggesting intra-bulbar modulation according to metabolic state. These results demonstrate that, akin to other animal models, hunger state affects OB activity in humans. Moreover, we show that the EBG method has the potential to measure internal metabolic states and, as such, could be used to study specificities in olfactory processing of individuals suffering from pathologies such as obesity or anorexia.

Phase-locked breathing does not affect episodic visual recognition memory but does shape its corresponding ERPs.

Recent studies have indicated that breathing shapes the underlying oscillatory brain activity critical for episodic memory, potentially impacting memory performance. However, the literature has presented conflicting results, with some studies suggesting that nasal inhalation enhances visual memory performance, while others have failed to observe any significant effects. Furthermore, the specific influence of breathing route (nasal vs. mouth) and the precise phase of the respiratory cycle during which stimuli are presented have remained elusive. To address this, we employed a visual recognition memory (VRM) and electroencephalography paradigm in which stimuli presentation was phase-locked to either inhalation or exhalation onset, using a within-subject design where participants performed the memory task while engaging in separate sessions of nose and mouth breathing. We show that neither breathing route nor breathing phase has a significant impact on VRM performance as measured by d-prime, with the data supporting the null hypothesis. However, we did find an effect of breathing phase on response bias, with participants adopting a more conservative decision criterion during exhalation. Moreover, we found that breathing phase during memory encoding shaped the late parietal effect (LPE) amplitude, while the Frontal Negative Component (FN400) and LPE during recognition were less impacted. While our study demonstrates that breathing does not shape VRM performance, it shows that it influences brain activity, reinforcing the importance of further research to elucidate the extent of respiratory influence on perception, cognition, and behavior.

Inconclusive evidence that breathing shapes pupil dynamics in humans: a systematic review.

More than 50 years ago, it was proposed that breathing shapes pupil dynamics. This widespread idea is also the general understanding currently. However, there has been no attempt at synthesizing the progress on this topic since. We therefore conducted a systematic review of the literature on how breathing affects pupil dynamics in humans. We assessed the effect of breathing phase, depth, rate, and route (nose/mouth). We followed the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines, and conducted a systematic search of the scientific literature databases MEDLINE, Web of Science, and PsycInfo in November 2021. Thirty-one studies were included in the final analyses, and their quality was assessed with QualSyst. The study findings were summarized in a descriptive manner, and the strength of the evidence for each parameter was estimated following the Grading of Recommendations Assessment, Development, and Evaluation (GRADE) approach. The effect of breathing phase on pupil dynamics was rated as "low" (6 studies). The effect of breathing depth and breathing rate (6 and 20 studies respectively) were rated as "very low". Breathing route was not investigated by any of the included studies. Overall, we show that there is, at best, inconclusive evidence for an effect of breathing on pupil dynamics in humans. Finally, we suggest some possible confounders to be considered, and outstanding questions that need to be addressed, to answer this fundamental question. Trial registration: This systematic review has been registered in the international prospective register of systematic reviews (PROSPERO) under the registration number: CRD42022285044.

Loss of olfactory sensitivity is an early and reliable marker for COVID-19.

Detection of early and reliable symptoms is important in relation to limiting the spread of an infectious disease. For COVID-19, the most specific symptom is either losing or experiencing reduced olfactory functions. Anecdotal evidence suggests that olfactory dysfunction is also one of the earlier symptoms of COVID-19, but objective measures supporting this notion are currently missing. To determine whether olfactory loss is an early sign of COVID-19, we assessed available longitudinal data from a web-based interface enabling individuals to test their sense of smell by rating the intensity of selected household odors. Individuals continuously used the interface to assess their olfactory functions and at each login, in addition to odor ratings, recorded their symptoms and results from potential COVID-19 test. A total of 205 COVID-19-positive individuals and 156 pseudo-randomly matched control individuals lacking positive test provided longitudinal data which enabled us to assess olfactory functions in relation to their test result date. We found that odor intensity ratings started to decline in the COVID-19 group as early as 6 days prior to the test result date (±1.4 days). Symptoms, such as sore throat, aches, and runny nose appear around the same point in time; however, with a lower predictability of a COVID-19 diagnosis. Our results suggest that olfactory sensitivity loss is an early symptom but does not appear before other related COVID-19 symptoms. Olfactory loss is, however, more predictive of a COVID-19 diagnosis than other early symptoms.

An olfactory self-test effectively screens for COVID-19.

Background: Key to curtailing the COVID-19 pandemic are wide-scale screening strategies. An ideal screen is one that would not rely on transporting, distributing, and collecting physical specimens. Given the olfactory impairment associated with COVID-19, we developed a perceptual measure of olfaction that relies on smelling household odorants and rating them online. Methods: Each participant was instructed to select 5 household items, and rate their perceived odor pleasantness and intensity using an online visual analogue scale. We used this data to assign an olfactory perceptual fingerprint, a value that reflects the perceived difference between odorants. We tested the performance of this real-time tool in a total of 13,484 participants (462 COVID-19 positive) from 134 countries who provided 178,820 perceptual ratings of 60 different household odorants. Results: We observe that olfactory ratings are indicative of COVID-19 status in a country, significantly correlating with national infection rates over time. More importantly, we observe indicative power at the individual level (79% sensitivity and 87% specificity). Critically, this olfactory screen remains effective in participants with COVID-19 but without symptoms, and in participants with symptoms but without COVID-19. Conclusions: The current odorant-based olfactory screen adds a component to online symptom-checkers, to potentially provide an added first line of defense that can help fight disease progression at the population level. The data derived from this tool may allow better understanding of the link between COVID-19 and olfaction.

The perception of odor pleasantness is shared across cultures.

Humans share sensory systems with a common anatomical blueprint, but individual sensory experience nevertheless varies. In olfaction, it is not known to what degree sensory perception, particularly the perception of odor pleasantness, is founded on universal principles,1-5 dictated by culture,6-13 or merely a matter of personal taste.6,8-10,12,14 To address this, we asked 225 individuals from 9 diverse nonwestern cultures-hunter-gatherer to urban dwelling-to rank the monomolecular odorants from most to least pleasant. Contrary to expectations, culture explained only 6% of the variance in pleasantness rankings, whereas individual variability or personal taste explained 54%. Importantly, there was substantial global consistency, with molecular identity explaining 41% of the variance in odor pleasantness rankings. Critically, these universal rankings were predicted by the physicochemical properties of out-of-sample molecules and out-of-sample pleasantness ratings given by a tenth group of western urban participants. Taken together, this shows human olfactory perception is strongly constrained by universal principles.

The human olfactory bulb processes odor valence representation and cues motor avoidance behavior.

Determining the valence of an odor to guide rapid approach-avoidance behavior is thought to be one of the core tasks of the olfactory system, and yet little is known of the initial neural mechanisms supporting this process or of its subsequent behavioral manifestation in humans. In two experiments, we measured the functional processing of odor valence perception in the human olfactory bulb (OB)-the first processing stage of the olfactory system-using a noninvasive method as well as assessed the subsequent motor avoidance response. We demonstrate that odor valence perception is associated with both gamma and beta activity in the human OB. Moreover, we show that negative, but not positive, odors initiate an early beta response in the OB, a response that is linked to a preparatory neural motor response in the motor cortex. Finally, in a separate experiment, we show that negative odors trigger a full-body motor avoidance response, manifested as a rapid leaning away from the odor, within the time period predicted by the OB results. Taken together, these results demonstrate that the human OB processes odor valence in a sequential manner in both the gamma and beta frequency bands and suggest that rapid processing of unpleasant odors in the OB might underlie rapid approach-avoidance decisions.

A non-invasive olfactory bulb measure dissociates Parkinson's patients from healthy controls and discloses disease duration.

Olfactory dysfunction is a prevalent non-motor symptom of Parkinson's disease (PD). This dysfunction is a result of neurodegeneration within the olfactory bulb (OB), the first processing area of the central olfactory system, and commonly precedes the characteristic motor symptoms in PD by several years. Functional measurements of the OB could therefore potentially be used as an early biomarker for PD. Here, we used a non-invasive method, so-called electrobulbogram (EBG), to measure OB function in PD and age-matched healthy controls to assess whether EBG measures can dissociate PDs from controls. We estimated the spectrogram of the EBG signal during exposure to odor in PD (n = 20) and age-matched controls (n = 18) as well as identified differentiating patterns of odor-related synchronization in the gamma, beta, and theta frequency bands. Moreover, we assessed if these PD-EBG components could dissociate PD from control as well as their relationship with PD characteristics. We identified six EBG components during the initial and later stages of odor processing which dissociated PD from controls with 90% sensitivity and 100% specificity with links to PD characteristics. These PD-EBG components were related to medication, disease duration, and severity, as well as clinical odor identification performance. These findings support using EBG as a tool to experimentally assess PD interventions, potentially aid diagnosis, and the potential development of EBG into an early biomarker for PD.

Acquired olfactory loss alters functional connectivity and morphology.

Removing function from a developed and functional sensory system is known to alter both cerebral morphology and functional connections. To date, a majority of studies assessing sensory-dependent plasticity have focused on effects from either early onset or long-term sensory loss and little is known how the recent sensory loss affects the human brain. With the aim of determining how recent sensory loss affects cerebral morphology and functional connectivity, we assessed differences between individuals with acquired olfactory loss (duration 7-36 months) and matched healthy controls in their grey matter volume, using multivariate pattern analyses, and functional connectivity, using dynamic connectivity analyses, within and from the olfactory cortex. Our results demonstrate that acquired olfactory loss is associated with altered grey matter volume in, among others, posterior piriform cortex, a core olfactory processing area, as well as the inferior frontal gyrus and angular gyrus. In addition, compared to controls, individuals with acquired anosmia displayed significantly stronger dynamic functional connectivity from the posterior piriform cortex to, among others, the angular gyrus, a known multisensory integration area. When assessing differences in dynamic functional connectivity from the angular gyrus, individuals with acquired anosmia had stronger connectivity from the angular gyrus to areas primary responsible for basic visual processing. These results demonstrate that recently acquired sensory loss is associated with both changed cerebral morphology within core olfactory areas and increase dynamic functional connectivity from olfactory cortex to cerebral areas processing multisensory integration.

Human sickness detection is not dependent on cultural experience.

Animals across phyla can detect early cues of infection in conspecifics, thereby reducing the risk of contamination. It is unknown, however, if humans can detect cues of sickness in people belonging to communities with whom they have limited or no experience. To test this, we presented Western faces photographed 2 h after the experimental induction of an acute immune response to one Western and five non-Western communities, including small-scale hunter-gatherer and large urban-dwelling communities. All communities could detect sick individuals. There were group differences in performance but Western participants, who observed faces from their own community, were not systematically better than all non-Western participants. At odds with the common belief that sickness detection of an out-group member should be biased to err on the side of caution, the majority of non-Western communities were unbiased. Our results show that subtle cues of a general immune response are recognized across cultures and may aid in detecting infectious threats.

Verbally Induced Olfactory Illusions Are Not Caused by Visual Processing: Evidence From Early and Late Blindness.

Olfactory perception is malleable and easily modulated by top-down processes such as those induced by visual and verbal information. A classic example of this is olfactory illusions where the perceived pleasantness of an odor is manipulated by the valence of a verbal label that is either visually or auditorily presented together with the odor. The mechanism behind this illusion is still unknown, and it is not clear if it is driven only by verbal information or if there is an interaction between language functions and visual mental imagery processes. One way to test this directly is to study early blind individuals who have little or no experience of visual information or visual mental imagery. Here, we did this by testing early blind, late blind, and sighted individuals in a classical paradigm where odors were presented with negative, neutral, and positive labels via speech. In contrast to our hypothesis-that the lack of visual imagery would render early blind individuals less susceptible to the olfactory illusion-early and late blind participants showed more amplified illusions than sighted. These findings demonstrate that the general mechanism underlying verbally induced olfactory illusions is not caused by visual processing and visual mental imagery per se.

Odor identity can be extracted from the reciprocal connectivity between olfactory bulb and piriform cortex in humans.

Neuronal oscillations route external and internal information across brain regions. In the olfactory system, the two central nodes-the olfactory bulb (OB) and the piriform cortex (PC)-communicate with each other via neural oscillations to shape the olfactory percept. Communication between these nodes have been well characterized in non-human animals but less is known about their role in the human olfactory system. Using a recently developed and validated EEG-based method to extract signals from the OB and PC sources, we show in healthy human participants that there is a bottom-up information flow from the OB to the PC in the beta and gamma frequency bands, while top-down information from the PC to the OB is facilitated by delta and theta oscillations. Importantly, we demonstrate that there was enough information to decipher odor identity above chance from the low gamma in the OB-PC oscillatory circuit as early as 100 ms after odor onset. These data further our understanding of the critical role of bidirectional information flow in human sensory systems to produce perception. However, future studies are needed to determine what specific odor information is extracted and communicated in the information exchange.

No Evidence That Hormonal Contraceptives Affect Chemosensory Perception.

The use of oral contraceptives (OC) in the form of a hormonal pill has been widespread for decades. Despite its popularity and long-time use, there is still much ambiguity and anecdotal reports about a range of potential side effects. Here, we addressed the potential effect of OC use on chemosensory perception. Previous research has almost exclusively focused on olfaction, but we expanded this to the trigeminal system and the sense of taste. We used Bayesian statistics to compare the olfactory, trigeminal, and taste detection abilities between a group of 34 normal cycling women and a group of 26 women using OC. Our results indicated that odor, trigeminal, and taste thresholds were not affected by the use of OC. Moreover, neither odor perception, nor taste perception was affected; all with Bayes factors consistently favoring the null hypothesis. The only exception to these results was odor identification where Bayes factors indicated inconclusive evidence. We conclude that effects of OC use on chemosensory perception are unlikely, and if present, likely are of no to little behavioral relevance.

Whole-brain modelling of resting state fMRI differentiates ADHD subtypes and facilitates stratified neuro-stimulation therapy.

Recent advances in non-linear computational and dynamical modelling have opened up the possibility to parametrize dynamic neural mechanisms that drive complex behavior. Importantly, building models of neuronal processes is of key importance to fully understand disorders of the brain as it may provide a quantitative platform that is capable of binding multiple neurophysiological processes to phenotype profiles. In this study, we apply a newly developed adaptive frequency-based model of whole-brain oscillations to resting-state fMRI data acquired from healthy controls and a cohort of attention deficit hyperactivity disorder (ADHD) subjects. As expected, we found that healthy control subjects differed from ADHD in terms of attractor dynamics. However, we also found a marked dichotomy in neural dynamics within the ADHD cohort. Next, we classified the ADHD group according to the level of distance of each individual's empirical network from the two model-based simulated networks. Critically, the model was mirrored in the empirical behavior data with the two ADHD subgroups displaying distinct behavioral phenotypes related to emotional instability (i.e., depression and hypomanic personality traits). Finally, we investigated the applicability and feasibility of our whole-brain model in a therapeutic setting by conducting in silico excitatory stimulations to parsimoniously mimic clinical neuro-stimulation paradigms in ADHD. We tested the effect of stimulating any individual brain region on the key network measures derived from the simulated brain network and its contribution in rectifying the brain dynamics to that of the healthy brain, separately for each ADHD subgroup. This showed that this was indeed possible for both subgroups. However, the current effect sizes were small suggesting that the stimulation protocol needs to be tailored at the individual level. These findings demonstrate the potential of this new modelling framework to unveil hidden neurophysiological profiles and establish tailored clinical interventions.

Wine experts' recognition of wine odors is not verbally mediated.

Experts have better memory for items within their domain of expertise. Critically, this does not depend on more efficient use of language. However, this conclusion is based mainly on findings from experts in visual and auditory domains. Olfactory experts constitute an interesting potential counterexample since language has been implicated to be critically involved in odor memory in previous studies. We examined the role language plays in odor recognition memory for wine experts, who typically display better wine odor memory than novices and who are also able to name odors better than lay people. This suggests wine experts' superior recognition memory for odors may be verbally mediated. In 2 experiments, recognition memory for wine odors, wine-related odors, and common odors was tested in wine experts and novices. The use of language was manipulated in Experiment 1 with an overt naming versus no-naming condition, and in Experiment 2, with a verbal interference task inhibiting covert verbalization. Across the two experiments the results showed wine experts have better recognition memory for wines, but not for wine-related or common odors, indicating their memory advantage is expertise specific. Critically, this effect was not verbally mediated, as there was no relationship between experts' ability to name wines and their memory for them. Likewise, directly inhibiting online use of verbalization did not affect memory for wine odors in experts. In sum, once expertise has been acquired, language does not play a causal role in recognition memory for odors. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Putting action into testing: Enacted retrieval benefits long-term retention more than covert retrieval.

Retrieval practice improves long-term retention. However, it is currently debated if this testing effect can be further enhanced by overtly producing recalled responses. We addressed this issue using a standard cued-recall testing-effect paradigm with verb-noun action phrases (e.g., water the plant) to prompt motor actions as a specifically powerful response format of recall. We then tested whether motorically performing the recalled verb targets (e.g., ?-the plant) during an initial recall test (enacted retrieval) led to better long-term retention than silently retrieving them (covert retrieval) or restudying the complete verb-noun phrases (restudy). The results demonstrated a direct testing effect, in that long-term retention was enhanced for covert retrieval practice compared to restudy practice. Critically, enactment during retrieval further improved long-term retention beyond the effect of covert memory retrieval, both in a congruent noun-cued recall test after 1 week (Experiment 1) and in an incongruent verb-cued recall test of nouns after 2 weeks (Experiment 2). This finding suggests that successful memory retrieval and ensuing enactment contribute to future memory performance in parts via different mechanisms.

Expertise Shapes Multimodal Imagery for Wine.

Although taste and smell seem hard to imagine, some people nevertheless report vivid imagery in these sensory modalities. We investigate whether experts are better able to imagine smells and tastes because they have learned the ability, or whether they are better imaginers in the first place, and so become experts. To test this, we first compared a group of wine experts to yoked novices using a battery of questionnaires. We show for the first time that experts report greater vividness of wine imagery, with no difference in vividness across sensory modalities. In contrast, novices had more vivid color imagery than taste or odor imagery for wines. Experts and novices did not differ on other vividness of imagery measures, suggesting a domain-specific effect of expertise. Critically, in a second study, we followed a group of students commencing a wine course and a group of matched control participants. Students and controls did not differ before the course, but after the wine course students reported more vivid wine imagery. We provide evidence that expertise improves imagery, exemplifying the extent of plasticity of cognition underlying the chemical senses.