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. 2020 Jul 30;15(7):e0236468. doi: 10.1371/journal.pone.0236468

A network model of affective odor perception

Yingxuan Liu 1, Alexander Toet 1,*, Tanja Krone 2, Robin van Stokkum 2, Sophia Eijsman 1, Jan B F van Erp 1,3
Editor: Alberto Greco4
PMCID: PMC7392242  PMID: 32730278

Abstract

The affective appraisal of odors is known to depend on their intensity (I), familiarity (F), detection threshold (T), and on the baseline affective state of the observer. However, the exact nature of these relations is still largely unknown. We therefore performed an observer experiment in which participants (N = 52) smelled 40 different odors (varying widely in hedonic valence) and reported the intensity, familiarity and their affective appraisal (valence and arousal: V and A) for each odor. Also, we measured the baseline affective state (valence and arousal: BV and BA) and odor detection threshold of the participants. Analyzing the results for pleasant and unpleasant odors separately, we obtained two models through network analysis. Several relations that have previously been reported in the literature also emerge in both models (the relations between F and I, F and V, I and A; I and V, BV and T). However, there are also relations that do not emerge (between BA and V, BV and I, and T and I) or that appear with a different polarity (the relation between F and A for pleasant odors). Intensity (I) has the largest impact on the affective appraisal of unpleasant odors, while F significantly contributes to the appraisal of pleasant odors. T is only affected by BV and has no effect on other variables. This study is a first step towards an integral study of the affective appraisal of odors through network analysis. Future studies should also include other factors that are known to influence odor appraisal, such as age, gender, personality, and culture.

1 Introduction

1.1 The affective appraisal of odors

Odors can effectively elicit affective responses [15], probably due to the high degree of overlap and connectivity between the neural systems mediating olfaction and emotion [610]. These affective responses mediate our perception of environmental input and can adapt our output, thus enabling us to respond in an appropriate way [11]. The affective response to odors is typically characterized by its valence (pleasantness or hedonic tone) and arousal [12, 13], while both dimensions are mediated by different neural substrates [14]. Brain imaging studies show that unpleasant and pleasant odors also activate different brain areas [8, 1519] in asymmetric ways [17, 20]. Unpleasant odors are processed faster than pleasant ones [17, 2123], eliciting specific patterns of autonomic [24, 25] and olfactomotor responses [26, 27] and specific neural activation [14, 16, 18, 20, 2830]. Also, unpleasant odors are also less prone to top-down influences such as priming [31], verbal context [32] and odor knowledge [33].

Pleasant odors positively affect mood and decrease arousal, while unpleasant odors have the opposite effect [34]. It has been observed that unpleasant odors increase skin conductance, heart rate [3537] and the startle reflex [3840] while pleasant odors decrease these parameters. As a result, odors can effectively be used to induce various emotional states [2, 4143] and desired behaviors [11]. In real-life settings, odors have for instance effectively been deployed to reduce patient stress in healthcare environments [4446], to influence shopping behavior in retail environments [47, 48] and to influence littering behavior in public environments [49]. Because the principal distinctive properties of food flavors are provided by olfaction rather than by taste cues [50], our culinary preferences are also to a large extent based on the affective appraisal of food odors. However, despite the important role of affect in olfaction, it is still largely unknown how affective appraisal and olfactory perception interact and converge in everyday life [9].

1.2 Factors related to the affective appraisal of odors

Factors that are known to be related to the affective appraisal of odors include odor sensitivity, odor intensity, odor familiarity (the feeling that an odor is known or has been perceived before: [33]) and core affective state [25, 5154]. Previous studies only investigated the correlations between specific subsets of these factors. As a result, the extent to which individual differences in these factors and their interrelations differentially influence the affective response of people to specific odors is still largely unknown [2]. In this section we will first present the available evidence for the mediating effects of sensitivity, familiarity, and core or baseline affective state on affective odor appraisal. Fig 1 represents the known relations between these different factors as hypothetical graphical network models for affective odor appraisal (hypothetical odor evoked affect or HOEA model). Given the aforementioned evidence for the existence of different processing channels for unpleasant and pleasant odors, we will distinguish between an unpleasant (UHOEA: Fig 1a) and a pleasant (PHOEA: Fig 1b) model. In the next sections, we will refer to the relations between the variables in both HOEA models (indicated by R1-R15 in Fig 1) to facilitate the discussion.

Fig 1. Hypothetical odor-evoked affect (HOEA) network models for (a) pleasant (PHOEA) and (b) unpleasant (UHOEA) odors.

Fig 1

These networks represent the relations that have been reported in the literature between different factors influencing the affective appraisal of odors. Yellow nodes: the valence (V) and arousal (A) components of the affective odor appraisal. Blue nodes: the observer’s baseline valence (BV) and arousal (BA) values. I: odor intensity. T: odor detection threshold. F: odor familiarity. Edge color represents the polarity of the partial correlations green = positive, red = negative, grey = positive for pleasant odors, negative for unpleasant odors. The edge labels serve to identify the relations for discussion in the text.

1.2.1 Intensity

Odor intensity is generally negatively correlated with valence (R1): the more intense being the more unpleasant [51, 55]. However, intensity and valence interact in complex ways [5558], involving both innately tuned and learned components. The polarity of the effect may also depend on the nature of the stimulus and on the perceiver’s personal characteristics [52, 56, 57, 59, 60]. As a result, several exceptions to R1 have been observed, with some odors showing a positive correlation between valence and intensity, some a negative, and others an inverted U-shape or even an absence of correlation [5557, 59]. Experience and learning significantly determine odor valence [61, 62]. Odor knowledge (identification) significantly enhances ratings of intensity, pleasantness and familiarity [63]. There are also indications that individuals with high detection thresholds may show a positive correlation of odor intensity with valence [56], although the evidence for this assumption is weak. Odor intensity is typically strongly positively correlated with subjective and autonomic indices of arousal (R2; [12]), independent of odor valence [14, 56, 57].

1.2.2 Familiarity

Familiarity is implicitly linked to the affective appreciation of our environment rather than to explicit source recognition [64]. Olfaction appears to serve novelty or change detection (possibly mediated by the amygdala: [9, 65, 66]), directing our attention to odors that are either unknown (not experienced before: categorical novelty) or do not fit our expectation or previous experience of a given situation (contextual novelty or misfit; see [64]).

Both direct and indirect effects of odor familiarity on affective odor appraisal have been reported in the literature.

The relation between familiarity and odor valence appears to be asymmetrical. For pleasant odors (Fig 1b), familiarity and odor valence are typically positively correlated: the more familiar an odor, the more pleasant it is judged (R3 in Fig 1b; e.g.: [12, 5153, 63, 6775]). For unpleasant odors, no consistent relation has been found (R3 is absent in Fig 1a) [25, 33, 51]. This finding agrees with the idea that different evaluative channels are involved in the processing of negatively and positively valenced stimuli [76]. In general, unpleasant odors are relatively less prone to top-down (cognitive) influences [3133]. A negativity bias for unidentifiable pleasant odors may for instance reflect a behavioral system designed for self-protection that elicits a warning or avoidance response when confronted with a positive but unfamiliar (unknown or unexpected: [64]) odor that may represent a potential health threat [77].

Familiarity and odor evoked arousal are negatively correlated, independent of odor valence (R4): the more familiar pleasant (e.g., the comforting and relaxing smell of a familiar environment or the perfume of a loved one) and unpleasant (e.g., the smells of smoke or decay, signaling threat or danger) odors are, the less arousing they are judged [25].

It also appears that familiarity can indirectly influence the affective appraisal of odors by modulating their intensity: participants perceive familiar odors as more intense than unfamiliar odors (R5; [51, 52, 63, 70]), which may ultimately influence their valence (R1; [63]). Note that familiarity may differentially affect the affective appraisal of an odor depending on a person’s history with it (e.g., due to a change in valence because of its contiguous presentation with a positive or negative event [78]).

1.2.3 Affective state

Both direct and indirect effects of affective state on the appraisal of affective stimuli have been reported in the literature.

Core or baseline affective state may have a direct impact on subsequent judgements through misattribution [7982]. People are inclined to make cognitive appraisals of unrelated topics and objects reflecting their core affective state (R6-R9; [81, 83]). In particular, they tend to attribute residual arousal from prior experiences to external cues in subsequent situations (R9; [84]). Since this may also be the case for the affective appraisal of odors, we hypothesize that BV and V (R6) and BA and A (R9) may be positively correlated (a carry-over effect), while BV and A (R7) and BA and V (R8) may be negatively correlated (a contrast effect).

Core affective state can also indirectly influence the affective appraisal of odors. Affective state modulates chemosensory event-related potentials [54] and affects odor intensity (R10, R11): it has been observed that emotions enhance odor intensity [85, 86], independent of odor valence [60, 87]. Furthermore, emotional valence also modulates the odor detection threshold: a negative emotional state reduces olfactory sensitivity (R12; [54, 86]. This may in turn influence odor associated affect by modulating the odor intensity: people with elevated thresholds perceive odors as being less intense (R13; [8891]). Although emotional arousal mediates the affective appraisal and intensity of odors [60, 86], there is currently no evidence that it directly modulates the odor detection threshold [86, 92].

1.3 Relation between valence and arousal

The general assumption of the independence between valence and arousal for the affective appraisal of affective stimuli has recently been questioned: although valence and arousal appear to be uncorrelated when valence is ambiguous, they tend to become correlated when valence is clear [9395]. Hence, these dimensions may be correlated (R14) for the affective appraisal of odors with a clear valence. For a wide range of different affective stimuli it has been found that arousal generally increases (a) with increasing valence for positively valenced stimuli and (b) with decreasing valence for negatively valenced stimuli [93]. Therefore, we assume that both variables are positively correlated for pleasant odors (R14 is positive in Fig 1b) and negatively correlated for unpleasant odors in the HOEA model (R14 is negative in Fig 1a).

Note that the valence of odors may change due to learning effects. While affective odors appraisal appears to be partly innate [9698], factors like the frequency and context of prior exposure, semantic knowledge, and cultural background can cause significant variations in hedonic perception between individuals and over the course of the human life-span [99]. For instance, odors that are initially perceived as neutral or positive may acquire a negative connotation and signal threat after they have been experienced in the context of negative life events [100].

1.4 Current study

The goal of this study was to explore the potential relations between the different variables in in our literature-based HOEA model (Fig 1). Since previous studies investigated these variables individually, there is currently no integral model for their interrelations. To fill this gap, we performed an observer experiment in which participants reported the valence and arousal, intensity and familiarity for a range of different odors, varying widely in hedonic valence. In addition, we measured the participants’ baseline affective state and detection threshold. We explored the relations between baseline affective state, odor familiarity, odor intensity and odor detection threshold, and their impact on affective odor appraisal through probabilistic network analysis [101104]. Network analysis is a data-driven exploratory approach to modelling, allowing model structure to spontaneously emerge from the statistical relationships among indicators, thereby eliminating the need to specify an a-priori model. Network analysis focusses on the direct relations between observed variables. Hence, network analysis and visualization can yield new insights into the relations between variables. In psychology, network analysis has recently become a popular alternative for latent variable modelling in exploratory studies of human behavior [101, 103, 105111]. Psychological networks consist of nodes representing observed variables (e.g., questionnaire items), connected by edges representing the statistical relationships between the variables (their pairwise interactions; [112]). Network analysis typically involves the following three steps [112]: (1) network estimation, (2) network analysis, and (3) network comparison.

In the rest of this paper we first present the methods, materials and techniques used in this study. Then we present the results and compare the network models that were estimated from our present results to the HOEA model that was based on findings from the literature. Finally, we discuss the implications of the current findings and the limitations of this study.

2 Methods

2.1 Participants

To conduct a power analysis (determine the adequate sample size) an expectation of the effect size is required. The network equivalent is an expected (weighted) network structure [112]. However, since this is the first study of its kind, no previous similar networks were available. Sample size was therefore determined from a general rule of thumb suggested in the literature; namely, three individuals per parameter [112]. Since the HOEA network has 14 edges, this means that this study required a minimal group size of 42 participants to meet this “rule of thumb”.

A total of 56 students (32 females, and 24 males, mean age = 24.3 years, SD = 4.6) from Utrecht University (Utrecht, the Netherlands) participated in this experiment. Participants were recruited through postings on social media and direct messaging. The exclusion criteria were age (younger than 18 years and older than 60 years), olfactory deficiencies (e.g., diseases, having a cold, smoking or drinking alcohol) and pregnancy. Participants were asked not to wear perfume, use deodorant or wear scented clothing on the testing day. All participants signed an informed consent form. The experimental protocol was reviewed and approved by the TNO Internal Review Board (TNO, the Netherlands: reference 2019–024) and was in accordance with the Helsinki Declaration of 1975, as revised in 2013 [113]. After completing the study, participants were offered a small compensation (5 Euro or study credits) for their participation.

2.2 Stimuli

In this study we measured odor-evoked valence and arousal for 40 different odors (see Table 1), ranging from unpleasant and arousing (e.g., feces, fish), via pleasant and calming (e.g., clove, cinnamon) to pleasant and stimulating (e.g., peach, caramel). To obtain a stimulus set with valence values distributed across the entire scale range, we complemented the revised 32-item “Sniffin’ Sticks” odor identification test, which contains neutral and pleasant smells (www.burghart-mt.de, see also: [114]), with eight additional odors that are typically perceived as unpleasant: burned wood, diesel fumes, dusty cave, metal, rhinoceros, tar (obtained from https://retroscent.com and indicated by the RS codes in Table 1) and with indole (unpleasant smell associated with feces) and wintergreen (typically perceived as less pleasant by Europeans: [34]; both obtained from www.hekserij.nl). The Sniffin’ Sticks identification test consists of two sets (a blue capped set and a purple capped set) of 16 numbered felt pens each, with tips that are impregnated with 4 mL of fluid odor substance. This test is normally used to assess an individual’s olfactory identification performance [115117]. We prepared eight extra sticks by injecting 4 mL of the additional unpleasant odor substances in empty Sniffin’ Sticks. Hence, our total stimulus set consisted of 40 sticks pens, numbered from 1 to 40 (see Table 1). Since extreme differences in intensity may confound the affective ratings because of the inverse valence-intensity relation, a panel consisting of three of the authors (SE, YL, AT) verified that the set of odor samples did not contain any outliers in intensity, prior to the experiments. To ensure the compatibility between the samples, we adopted the criterion set by the developers of the Sniffin’ Sticks that all intensities should be within about 25% of the mean intensity [116]. The same set of sticks was used during the entire experiment. All samples were prepared in compliance with the safety Standards of the International Fragrance Association [118].

Table 1. Mean (SD) valence, arousal, familiarity and intensity ratings for all odors used as stimuli in this study.

The Sniffin’ Sticks B and P codes refer to the Blue and Purple identification test sets (www.burghart-mt.de). The RS codes refer to the RetroScent product code (https://retroscent.com). Odors with a negative mean valence rating are printed in boldface.

ID Label Code Valence Arousal Familiarity Intensity
1 Anise Sniffin’ B15 0.34 (2.09) 0.05 (2.02) 72.63 (23.39) 54.34 (20.91)
2 Apple Sniffin’ B11 0.84 (2.08) 0.20 (1.98) 55.07 (23.54) 61.89 (19.19)
3 Banana Sniffin’ B5 1.47 (1.72) 0.48 (2.24) 77.25 (17.76) 60.18 (20.14)
4 Burned wood RS-420 -1.54 (2.15) 0.76 (2.09) 49.86 (28.08) 73.07 (21.13)
5 Caramel Sniffin’ P15 2.25 (1.39) 1.11 (2.25) 80.50 (13.62) 58.75 (20.44)
6 Cinnamon Sniffin’ B3 0.98 (1.98) 0.22 (2.40) 61.88 (29.39) 54.25 (23.22)
7 Cloves Sniffin’ B12 -0.33 (2.30) -0.02 (2.23) 51.38 (29.16) 62.88 (21.05)
8 Coconut Sniffin’ P9 1.94 (1.59) 0.87 (2.10) 77.32 (19.03) 56.16 (19.42)
9 Coffee Sniffin’ B10 0.77 (2.26) 0.33 (2.23) 72.52 (27.13) 58.75 (22.12)
10 Coke Sniffin’ P2 0.37 (1.77) -0.48 (1.86) 48.11 (25.65) 49.84 (20.35)
11 Diesel fumes RS-423 -1.76 (1.55) -0.04 (2.22) 48.84 (23.97) 60.86 (21.47)
12 Dusty cave RS-425 -0.20 (1.82) -0.76 (1.89) 36.59 (22.94) 43.00 (25.00)
13 Eucalyptus Sniffin’ P7 0.66 (2.09) 0.21 (2.11) 73.55 (22.77) 70.36 (19.14)
14 Fish Sniffin’ B16 -2.07 (1.74) 0.84 (2.34) 63.16 (27.69) 71.39 (26.26)
15 Garlic Sniffin’ B9 -1.18 (2.20) 0.79 (2.32) 68.30 (26.61) 77.21 (18.14)
16 Ginger Sniffin’ P8 0.01 (1.91) -0.52 (1.87) 47.38 (24.41) 55.36 (20.81)
17 Grapefruit Sniffin’ P4 0.83 (1.91) 0.12 (2.11) 53.20 (24.58) 55.48 (19.65)
18 Grass Sniffin’ P5 0.07 (2.04) -0.14 (2.03) 67.43 (22.45) 63.93 (21.36)
19 Feces Indole -1.97 (1.86) 0.53 (2.16) 40.61 (24.86) 65.13 (22.17)
20 Lavender Sniffin’ P10 0.93 (1.88) -0.08 (2.20) 66.91 (25.23) 56.64 (18.45)
21 Leather Sniffin’ B2 -0.57 (1.91) -0.71 (1.54) 41.52 (25.13) 46.16 (20.95)
22 Lemon Sniffin’ B6 1.49 (1.73) 0.48 (2.12) 60.14 (25.38) 53.70 (21.15)
23 Lilac Sniffin’ P3 0.92 (2.02) -0.32 (2.05) 68.00 (22.94) 59.23 (20.04)
24 Liquorice Sniffin’ B7 0.92 (2.13) 0.00 (1.93) 78.23 (24.00) 57.04 (19.83)
25 Melon Sniffin’ P11 1.58 (1.68) -0.12 (2.38) 66.66 (21.52) 54.64 (22.07)
26 Metal RS-426 -0.71 (1.75) -0.59 (1.82) 36.95 (25.43) 50.09 (22.73)
27 Mushroom Sniffin’ P13 -1.28 (2.10) 0.51 (2.12) 45.13 (29.10) 66.30 (18.68)
28 Onion Sniffin’ P16 -1.73 (2.03) 1.00 (2.12) 55.66 (28.30) 69.20 (21.28)
29 Orange Sniffin’ B1 2.04 (1.30) 0.57 (2.23) 76.25 (19.69) 54.88 (24.05)
30 Peach Sniffin’ P12 2.51 (1.67) 1.55 (2.07) 79.70 (19.67) 61.84 (22.93)
31 Pear Sniffin’ P1 1.14 (1.74) 0.02 (2.11) 59.00 (22.74) 57.23 (19.97)
32 Peppermint Sniffin’ B4 1.63 (1.80) 0.57 (2.02) 84.84 (16.30) 64.80 (23.05)
33 Pineapple Sniffin’ B13 1.21 (2.07) 0.22 (2.17) 58.93 (26.68) 64.77 (18.26)
34 Raspberry Sniffin’ P6 2.20 (1.23) 0.57 (2.18) 66.54 (18.73) 54.77 (21.88)
35 Rhinoceros RS-424 -2.10 (1.48) 0.46 (2.31) 49.34 (25.58) 70.84 (20.02)
36 Rose Sniffin’ B14 1.43 (1.77) -0.09 (2.13) 72.77 (16.78) 61.23 (21.94)
37 Smoked meat Sniffin’ P14 -0.92 (1.82) -0.57 (1.92) 44.16 (26.65) 58.54 (21.82)
38 Tar RS-401 -1.23 (2.20) 0.63 (2.07) 52.71 (26.88) 72.59 (20.06)
39 Turpentine Sniffin’ B8 -0.87 (1.94) -0.55 (2.01) 46.91 (26.11) 57.77 (18.90)
40 Wintergreen Gaultheria oil -0.99 (2.19) 0.02 (2.02) 56.48 (29.89) 69.30 (21.12)

2.3 Measures

2.3.1 Odor detection threshold

Odor detection thresholds were measured using the standard “Sniffin’ Sticks” odor threshold test (www.burghart-mt.de) in combination with a single-staircase, triple-forced-choice procedure [117]. The test comprises 16 triplets of pens (total of 48 pens). The three pens in each triplet are distinguished by the color of their cap (red, green and blue). Red pens are impregnated with phenylethylalcohol (PEA) diluted in a solvent according to decreasing concentrations. Blue and green pens are only impregnated with solvent. During the test, participants were blindfolded with a sleep mask to prevent them from recognizing the odorant-containing pens. For odor presentation, a pen’s cap was removed by the experimenter for about 3 s and the pen’s tip was placed approximately 2 cm below both nostrils of the participant. The three pens of a triplet (two containing only the solvent and one containing also the odorant) were presented in a randomized order. Participants were asked to detect the odor-containing pen in each triplet (forced choice). Triplets were presented at intervals of approximately 20 s. Reversal of the staircase toward lower concentrations was triggered either when the odor was correctly detected in two successive trials or toward higher concentrations when the odor was not detected in a trial. The total number of reversals was seven, and the threshold (T) was defined as the arithmetic mean of the last four staircase reversals. There was no absolute number of correct responses required. The subjects’ scores ranged between 1 (lowest sensitivity or highest threshold: no odor detected) and 16 (highest sensitivity or lowest threshold).

2.3.2 Valence and arousal

The graphical EmojiGrid affective self-reporting tool (Fig 2; [119]) was used to measure subjective valence and arousal. The EmojiGrid is a Cartesian axes system similar to the Affect Grid [120], but the verbal labels on the midpoints and endpoints of the axes are replaced with emoji showing facial expressions. Also, additional emoji are inserted between the midpoints and the endpoints of each axis (resulting in five emoji on each side of the grid), and one (neutral) emoji is placed in the center of the grid, resulting in a total of 17 emoji on the grid. The central emoji with a neutral expression serves as a baseline or anchor point. The facial expressions of the emoji vary from disliking (unpleasant) via neutral to liking (pleasant) along the horizontal (valence) axis, and gradually increase in intensity along the vertical (arousal) axis. The facial expressions are defined by the eyebrows, eyes and mouth configuration of the face, and are inspired by the Facial Action Coding System [121]. The arousal dimension is represented by the opening of the mouth and the shape of the eyes, while the valence dimension is represented by the concavity of the mouth, the orientation and curvature of the eyebrows, and the vertical position of these features in the face area (representing a slightly downward looking face for lower arousal values and a slightly upward looking face for higher valence values). Users respond by clicking on a point inside the grid that best represents their affective appraisal of the stimulus.

Fig 2. The EmojiGrid: An emoji labeled Affect Grid for the measurement of odor-related affective associations.

Fig 2

The facial expressions of the emoji vary from disliking unpleasant via neutral to liking pleasant along the horizontal valence axis, and gradually increase in intensity along the vertical arousal axis.

At the start of the experiment participants first rated their baseline affective state on the EmojiGrid. In the rest of the experiment they used the EmojiGrid to rate their affective appraisal of the 40 different odor stimuli. All ratings were scaled to a range between -4 and 4.

2.3.3 Odor intensity and familiarity

Familiarity (F) and intensity (I) of each odor were measured with two single-item questions: “How intense do you perceive the scent?” and “How familiar are you with the scent?”. Participants rated these items by dragging a slider under each question to a value between 0 and 100. The slider defaulted at 50. Participants could see a tooltip with the current slider value while rating.

2.4 Procedure

The tests were performed in a quiet, well-ventilated room to avoid the presence of any residual odors. The experimenter wore odorless cotton gloves during the entire experiment. A computer was used to register all responses and to suggest a random stimulus presentation order to the experimenter.

Fig 3 shows the timeline of the events in the experimental procedure.

Fig 3. Timeline of the events in the experimental procedure.

Fig 3

Upon their arrival at the laboratory, the participants were welcomed by the experimenter and received a verbal introduction and instructions. Then, they read and signed an informed consent. Next, they filled in their nationality, age and gender. Then the EmojiGrid was presented on a computer screen and the participants were asked to study it carefully. They were informed that they could respond by clicking on a point inside the grid that best represented their emotional state.

The experiment consisted of two parts. In the first part the odor detection threshold of the participants was determined. In the second part the participants rated the intensity, familiarity and the subjective valence and arousal for each of the 40 different odors. Before starting the odor measurements, the participants first rated their momentary affective state (valence and arousal) using the computer-based EmojiGrid. Then they were blindfolded, and the odor detection threshold test started. After finishing the threshold test, the participants took off their blindfolds and the experimenter started the odor appraisal test. The participants were explicitly asked not to attempt to identify the smells since knowledge of odor sources may influence their valence, intensity and familiarity [33, 51, 63, 73, 122]. During the experiment, the experimenter presented each of the 40 scent pens once (after removing the cap of the pen) for about 5 seconds at a distance of about 2 cm from the edge of both nostrils of the participant. The presentation order was randomized over the participants. The participants sniffed following a brief verbal command (natural sniffing is known to provide optimal odor perception: [123]). Immediately after sniffing the pen was removed (and its cap replaced by the experimenter), and the participants were given at least 30 s to smell fresh air (to reduce potential effects of olfactory adaptation and habituation: [124]). During this interval, participants rated their affective appraisal (valence and arousal), intensity and familiarity of the smell. The entire experiment lasted about an hour.

2.5 Data analysis

2.5.1 General statistics

IBM SPSS Statistics 25 (www.ibm.com) was used to inspect the data for outliers (standardizing all ratings of intensity, familiarity, valence and arousal for each odor and for each participant) and to compute the mean values for valence and arousal for each odor over all participants.

Matlab 2019a (www.mathworks.com) was used to investigate the relation between the (mean) valence and arousal ratings and plot the data. The Curve Fitting Toolbox (version 3.5.7) in Matlab was used to compute a least-squares fit of a quadratic function to the data points. Based on this analysis (the mean valence ratings) the odors in the stimulus set were classified as either positive or negative.

All further data analysis was done in R version 3.6.0 (R Core Team, www.r-project.org) in R Studio 1.2.1335 (www.rstudio.com). The exact version numbers of all R packages used are documented in the S1 Data.

In all statistical analyses, a probability level of p < .05 was considered as statistically significant. To attenuate interindividual variance (as this is not the main interest of this paper) while retaining within-subject variance, we converted the valence, arousal, intensity and familiarity scores per individual and per odor valence set (pleasant/unpleasant) to z-scores [125]. Participants with standardized values exceeding two standard deviations from the mean were considered as outliers.

2.5.2 Network estimation

The most popular method to estimate network models for continuous and normally distributed data is the Gaussian Graphical Model (GGM: [126]). The GGM estimates a network of regularized partial correlations, thereby controlling for spurious relationships. When continuous data are not normally distributed, a transformation should be applied (e.g., a nonparanormal transformation; [127], see also [104]) to Gaussianize the input before estimating the GGM. In the resulting network, two connected variables are dependent after controlling for all other variables in the dataset. Thus, an edge connecting two nodes represents their conditional dependence given all other nodes. The absence of an edge between two nodes indicates that they are conditionally independent given all other nodes. The GGM has extensively been applied to psychological data [106, 107, 110, 128].

In this study we constructed GGMs to investigate the relations between affective state, odor sensitivity, odor intensity, odor familiarity and the affective appraisal of (positive and negative) odors. We used the nonparanormal transformation huge.npn from the huge R package [129] to normalize the data. Note that partial correlations can differ from zero due to sampling variation and may therefore represent false relations [130]. We therefore regularized our models with the graphical LASSO (Least Absolute Shrinkage and Selection Operator: [131]) algorithm, using the R packages glasso [131] and qgraph [132]. This procedure eliminates weak edges and returns a sparse network by driving low values of partial correlations to zero [104]. A sparse network is a parsimonious one that best accounts for the covariance among nodes while minimizing the number of edges. The LASSO algorithm first generates 1000 different network models with different degrees of sparsity (ranging from fully connected to fully disconnected), for 1000 different values of the tuning parameter λ that controls the level of sparsity [104, 133]. Then, it selects the model with the minimal EBIC (Extended Bayesian Information Criterion: [134]) value, given a value of the hyperparameter γ (which controls the trade-off between including potentially true edges and eliminating potentially false edges: [112]). The hyperparameter γ is usually set between zero and 0.5 [112]. As the value of γ approaches 0.5, the EBIC will favor a simpler network with fewer edges. In this study we set γ to its recommended default value of 0.5 [128, 135] to maximize the likelihood that the edges in the resulting network models represent genuine relations. Estimating a GGM with the glasso algorithm in combination with the EBIC model selection has been shown to reliably retrieve the true network structure [135] and is currently the dominant method for estimating GGMs in psychological network estimation [103, 104, 112].

The networks were visualized with the R package qgraph [132]. The node locations were determined using a modified version of the Fruchterman–Reingold algorithm [136, 137] for weighted networks [132], to ensure that strongly connected nodes with many edges in common are placed close to one another.

2.5.3 Network analysis

Once a network has been computed, different methods can be used to analyze its structure. Visual inspection is a useful first step that provides relevant information with minimal effort, especially for small networks [138, 139]. A more formal analysis of the relative importance of nodes in a network can for instance be performed by quantifying their direct (strength centrality) or indirect (closeness centrality) connectivity with other nodes or their mediating capacity between other nodes (betweenness centrality). When two networks need to be compared, their layout should be constrained to allow visual comparison (e.g. by using the averageLayout option in the qgraph R package: [132]) and permutation tests can be used quantify their structural similarity (e.g., by using the R package NetworkComparisonTest: [140]). Each of these steps will be discussed in the next sections.

2.5.3.1 Centrality indices. The importance of an individual node in a network is reflected in the number and strength of its connections to other nodes. In network analysis this is generally operationalized through three centrality indices: node strength (quantifying how strongly a node is directly connected to other nodes), closeness (quantifying how strongly a node is indirectly connected to other nodes), and betweenness (the number of times a node lies on the shortest path between two other nodes [130, 141]). To investigate the extent to which the individual variables (nodes) in our models play a mediating role in odor-evoked affect, we used the centrality_auto function to compute their strength, closeness and betweenness indices and we visualized the results (as z-scores to ensure comparability between networks) with the centralityPlot function, both from the R package qgraph [132]. Node strength is computed as the sum of the absolute weights of all edges connected to a node. A strength-central node is one that strongly affects other nodes. The closeness centrality of a node indicates the average distance from all other nodes in the network and is computed as the inverse of the sum of the shortest distances between the node and all other nodes. A closeness-central node is affected strongly (either directly or indirectly) by other nodes in a network. The betweenness centrality of a node is computed as the number of times that the node is on the shortest path between any two other nodes. A betweenness-central node connects a large number of other nodes, serving a bridge function. We quantified the stability of the centrality indices by their correlation stability (CS) coefficient, the value of which should preferentially exceed 0.5 [112].

2.5.3.2 Accuracy and stability. We used the R package bootnet [112] to evaluate the robustness (in terms of accuracy and stability) of the estimated networks through a nonparametric bootstrap sampling procedure [142].

First, we assessed the accuracy of the edge weights by computing and plotting the 95% confidence intervals (CIs) for each edge from a distribution of edge weights generated by sampling the data 1,000 times with replacement [107, 112, 143].

Next, we evaluated the stability of the networks by repeatedly correlating the centrality indices of the original data with the centrality indices calculated from subsamples comprising progressively fewer cases. The number of bootstraps was again set to 1,000. A centrality index is considered less stable when its correlation value decreases with a reduction of the sample size. This is quantified by the correlation stability coefficient (CS-coefficient), which represents the maximum proportion of cases that can be dropped while maintaining 95% probability that the correlation between the centrality index of the full dataset and that of the subset is at least .70 (denoted as CS(cor = .70); the value of .7 was chosen since this is typically regarded as a large effect: [144]). CS-coefficients above .5 indicate a high stability, while a minimum CS-coefficient of .25 is recommended for sufficient stability to warrant further interpretation of the centrality indices [107, 112].

2.5.4 Network comparison

We compared the structure of the unpleasant (UOEA) and pleasant (POEA) odor evoked affect networks in several ways.

First, we performed a visual comparison between the UOEA and POEA networks. Then, we computed a similarity index by correlating the edge weights across the two networks (i.e., by correlating their regularized partial correlation matrices: [145]). This index measures the correspondence between the strength of the network connections in both models. If the correlation equals one, the connections in both networks are perfectly linearly related, meaning that both networks essentially have the same structure; if it equals zero, the networks have no detectable linear correspondence; if it equals minus one, the networks are exact opposites.

Next, we formally tested their difference using the R package NCT (Network Comparison Test: [146]). The NCT is a two-tailed permutation test in which the difference between two groups is calculated repeatedly (10,000 times) for randomly regrouped cases. This produces a distribution of values under the null hypothesis (i.e., assuming equality between the groups) enabling one to test whether the observed difference in global strength differs significantly (p < .05) between two networks. The NCT can test invariance of structure and invariance of global strength. Invariance of structure is tested by comparing the largest observed difference (M) between corresponding edges in the two networks to that observed under permutation. Invariance of global strength (S) is tested by comparing the value of this parameter to that observed under permutation. Previous network research has shown that strength is typically the most stable and reliable centrality index [128, 143].

Results

Four participants were identified as outliers (their standardized ratings exceeded two standard deviations from the mean). Two of them gave an extremely low valence rating for the peach odor (ID = 31, see Table 1). Two other participants gave extremely low ratings for either the intensity of the pineapple odor (ID = 21) and the familiarity of the peppermint odor (ID = 2). After excluding these four participants from further analysis the remaining sample consisted of 52 participants (31 females and 21 males, with a mean age of 24.3 years, SD = 4.7).

First, we determined the mean ratings for valence, arousal, familiarity and intensity for each odor over all participants. The results are listed in Table 1. Fig 4 shows that the overall relation between mean valence and arousal can be described by a U-shaped (quadratic) form: odors scoring near neutral (zero) on mean valence have the lowest mean arousal ratings, while odors scoring either high (pleasant) or low (unpleasant) on mean valence show higher mean arousal ratings. Hence, odors with opposite mean valence ratings may yield similar mean arousal ratings. Because of the functional dichotomy that may exist in the relation between valence and the other variables that are measured in this study (e.g., the relation between valence and familiarity: [25]), we separately analyzed the results for unpleasant (odors with negative mean valence ratings) and pleasant (odors with positive mean valence ratings) stimuli. We classified the 16 odors with mean valence ratings below neutral as unpleasant stimuli (the odors with ID: 4, 7, 11, 12, 14, 15, 19, 21, 26, 27, 28, 35, 37, 38, 39, 40; see Table 1 and Fig 4) and the 24 odors with mean valence ratings above neutral as pleasant stimuli (the odors with ID: 1, 2, 3, 5, 6, 8, 9, 10, 13, 16, 17, 18, 20, 22, 23, 24, 25, 29, 30, 31, 32, 33, 34, 36). The mean intensity ratings listed in Table 1 show that the set of odor stimuli contained no outliers in intensity: all intensities were within about 28% of the mean intensity (which closely agrees with the criterion of 25% set by the developers of the Sniffin’ Sticks [116]).

Fig 4. Relation between mean valence and arousal ratings for the odors used in this study.

Fig 4

The numbers correspond to the identifiers in Table 1. Red numbers correspond to odors that received a negative mean valence rating, while blue numbers indicate odors that received a positive mean valence rating. The gray curve represents a quadratic fit to data points (R2 = .59).

Next, we estimated two network models: one for pleasant odors and one for unpleasant odors. In the following we will use the previously introduced abbreviations for the variable names (see Fig 1) to designate each node in these networks: BV and BA indicate respectively the valence and arousal components of the participant’s baseline affective state (measured at the start of the experiment), T designates the detection threshold, F represents the familiarity of an odor, I its intensity, while V and A represent respectively the valence and arousal associated with an odor. Unpleasant and pleasant odor-evoked affect will be referred to as UOEA and POEA respectively.

3.1 Network estimation

Fig 5 shows a graphical representation of the estimated (regularized) partial correlation network models (Gaussian Graphical Models) for (Fig 5a) unpleasant odor evoked affect (UOEA) and (Fig 5b) pleasant odor evoked affect (POEA), based on the sample of 52 participants that evaluated 16 unpleasant and 24 pleasant odors. Table 2 lists the partial correlations between the different variables in both networks. The resulting network structures are parsimonious due to the LASSO estimation: the UOEA and POEA networks (each containing 7 nodes) respectively have only 6 (3 positive and 3 negative) and 7 (5 positive and 2 negative) non-zero edges out of the 21 (= 6*7/2) possible edges.

Fig 5. Estimated partial correlation networks for unpleasant (a) and pleasant (b) odor evoked affect, based on a sample of 52 participants that evaluated 16 unpleasant and 24 pleasant odors.

Fig 5

Nodes represent the observed variables (for the meaning of the node labels we refer to the text), while green and red edges represent positive and negative partial correlations. Edge labels represent the relational identifiers from Fig 1. The width of the edges increases with the magnitude of the correlations and is scaled to the strongest edge (and therefore not comparable between graphs in an absolute sense).

Table 2. Partial correlations between the different variables in the estimated UOEA and POEA networks (Fig 5).

Dashes represent relations from the HOEA model that do not appear in the estimated networks.

Label Relation UOEA POEA
R1 I − V -0.36 -0.11
R2 I − A 0.25 0.24
R3 F − V 0.32 0.40
R4 F − A - 0.12
R5 F − I 0.31 0.27
R6 BV − V - -
R7 BV − A - -
R8 BA − V - -
R9 BA − A - -
R10 BV − I - -
R11 BA − I - -
R12 BV − T -0.42 -0.42
R13 T − I - -
R14 V − A -0.18 0.21

3.2 Network analysis

3.2.1 Centrality indices

Both resulting (UOEA and POEA) networks consist of three independent (unconnected) components: a trivial graph consisting of one node (BA), a simple graph consisting of two nodes (BV and T), and a connected graph consisting of the remaining four nodes (A, F, I and V). Table 3 lists the three (standardized, z-scored) centrality indices (strength, betweenness and closeness) for the nodes in the largest (4-node connected component of both the UOEA (Fig 5a) and POEA (Fig 5b) network models. It appears that these nodes differ substantially in their centrality estimates.

Table 3. Strength, closeness and betweenness centrality indices (using standardized z-scores to facilitate interpretation) for each node in the estimated networks (see Fig 5) for unpleasant and pleasant odor evoked affect, estimated from 1,000 bootstrap replications with the adaptive LASSO algorithm. Maximal values are printed in boldface.
Unpleasant odors Pleasant odors
Node Betweenness Closeness Strength Betweenness Closeness Strength
A -0.38 -1.17 -0.29 -0.59 -1.43 0.25
F -0.38 -0.39 0.42 1.46 0.68 1.09
I 2.27 1.14 1.27 -0.59 0.07 0.43
V -0.38 0.42 1.07 1.46 0.68 0.83

I is the most central node in the UOEA network, with the highest scores on all three centrality indices. This implies that I most significantly (directly and indirectly) contributes to the affective appraisal of unpleasant odors. F scores highest on all three centrality indices in the POEA network, indicating that this factor significantly contributes to the affective appraisal of pleasant odors.

3.2.2 Accuracy and stability

The accuracy and stability of the centrality indices was investigated by a case-dropping bootstrapped sampling procedure with 1,000 samples.

Fig 6 shows the bootstrapped 95% confidence intervals for the edge-weights in the estimated UOEA and POEA networks. This figure shows that the confidence intervals of positive and negative edges do not overlap, meaning that edges with opposite signs in Fig 5 are significantly different. However, the edge weights of some positive edges in Fig 5 (e.g., R2, R5, R14) may not be significantly different on the population level, since their confidence intervals show a large degree of overlap.

Fig 6. Bootstrapped 95% confidence intervals gray areas for the edge-weights in the estimated networks for unpleasant (A) and pleasant (B) odor evoked affect.

Fig 6

The red line connects the sample values, the black line the bootstrap means. The gray area represents the CIs. Each point represents one edge in the network, ordered from the edge with the highest weight (top) to the edge with the lowest weight (bottom). The labels along the outside of the vertical axis indicate the relations between different variable pairs (see text for the abbreviations) while the corresponding labels on the inside correspond to the edge labels relation numbers in the hypothetical odor evoked affect HOEA model (see Fig 1).

Fig 7 shows the stability plots for the centrality indices strength and betweenness (note that closeness could not be evaluated because of the infinite distance between the unconnected components) for both the unpleasant (Fig 7a) and pleasant (Fig 7b) odor evoked affect network models from Fig 5. These figures show that node strength (the associations of a node with its immediate neighbors) is highly stable for variations in sample size in both networks. Betweenness (connecting other nodes) shows a somewhat steeper decrease in accuracy with sample size than strength, especially in the POEA network. As a result, we cannot confidently conclude that any node in the POEA network is significantly more central than any other.

Fig 7. Stability of the central indices strength and betweenness of the estimated networks for unpleasant (A) and pleasant (B) odor evoked affect.

Fig 7

Data points represent the average correlation between the estimates based on subsamples, expressed as a percentage of original number of cases and the entire original sample. Areas indicate the range between the 2.5th and 97.5th quantiles.

Table 4 lists the correlation stability coefficients for the network centrality indices. Strength centrality is the only stable network characteristic. For both networks (UPOEA and POEA), the strength stability coefficient is 0.75, exceeding the recommended minimum value of 0.25. This means that strength centrality induces a meaningful order on the nodes in the networks.

Table 4. Correlation stability coefficients for the network centrality indices.

The CScor = 0.7 coefficients represent the maximum proportion of cases that can be dropped such that the correlation between the original centrality metric and those of the sampled subsets is 0.7 or higher with 95% probability.

Network model Strength Closeness Betweenness
UOEA 0.75 0.00 0.13
POEA 0.75 0.00 0.13

3.3 Network comparison

3.3.1 Global network structure

A visual comparison of the UOEA and POEA networks in Fig 5 shows that their structure is very similar: six edges occur in both networks (R1, R2, R3, R5, R12, R14). This observation is confirmed by the Pearson correlation between the adjacency matrices of both networks, which is r = .81, indicating a high degree of similarity. Also, the NCT revealed that the global strength of the UOEA network (1.85) does not differ significantly from that of the POEA network (1.75; p = .56). Corresponding relations in both networks have similar absolute partial correlation strengths (see Table 2) and the same polarity, except for the relation R14: as expected (see section 1.3) valence and arousal are positively correlated for positive odors and negatively correlated for unpleasant odors.

The main difference between both network structures is the relation between F and A (R4): familiarity only appears to (positively) influence the arousing quality of positive odors, but not of negative odors.

3.3.2 Intensity

In both emerging network models, odor intensity is negatively correlated with valence in (R1): the more intense being the more unpleasant. However, the relation is stronger for unpleasant odors than for pleasant odors. Odor intensity is positively correlated with subjective arousal (R2): odors that are more intense are rated as more arousing, independent of odor valence. These results both agree with the general findings in the literature, as embodied in both HOEA models (Fig 1).

3.3.3 Familiarity

Familiarity and valence are strongly positively correlated (R3): the more familiar an odor, the more pleasant it is judged, independent of odor valence. Familiarity is weakly positively correlated with odor evoked arousal for pleasant odors (R4), while no relation emerges for unpleasant odors. This is in contrast with the HOEA model, that predicts a negative correlation between odor familiarity and odor evoked arousal, independent of odor valence. In agreement with the HOEA model, familiarity is positively correlated with odor intensity in both networks (R5): participants perceive familiar odors as more intense than unfamiliar odors, independent of odor valence.

3.3.4 Affective state

The current results show no modulating effect of the arousal and valence components of the observer’s baseline affective state on both the intensity and the affective appraisal of odors, independent of their valence (relations R6-R11 from the HOEA model are absent in the UOEA and POEA models).

In agreement with the HOEA model, the valence component of baseline affective state (BV) correlates strongly and negatively with the odor detection threshold (R12). This agrees with the finding that negative mental states reduce odor sensitivity [54, 86], whereas a positive mental state can enhance odors sensitivity [86]. Contrary to our expectations, we find no negative correlation between T and I (R13).

4 Discussion

Based on a review of the literature we identified four individual factors that can influence odor-evoked affect as measured in term of valence and arousal: baseline affective state, odor sensitivity (detection threshold), odor intensity and odor familiarity. However, the exact nature of the relations between each of these variables and their influence on odor evoked affect are still largely unknown. To investigate these relations, we first constructed a hypothetical relational model, based on the small amount of literature that is currently available. Then we performed an observer experiment to collect data that can be used to verify this model. We used network analysis to explore the relations between the measured variables and odor-evoked affect through (regularized) partial correlations. This technique offers a data-driven view of the salient relationships between the variables of interest: relations emerge as partial correlations between the individual variables. Since the resulting GGMs are undirected networks it is not possible to discern causal relations. However, the absence of an edge between two factors in these models provides strong evidence that neither factor causes the other.

Because of the functional dichotomy that may exist in the relation between valence and the other variables that are measured in this study (e.g., the relation between valence and familiarity: [25]), we investigated the results for unpleasant (odors with negative mean valence ratings) and pleasant odors (odors with positive mean valence ratings) separately. Hence, we obtained two models: one for unpleasant odor evoked affect (UOEA model) and one for pleasant odor evoked affect (POEA model). It appears that both models are highly similar. The positive correlations between F and V for pleasant odors [12, 5153, 6773], and between F and I [52, 70], and between I and A [12, 14, 56, 57] that have been reported in the literature, also emerge in both networks. Other known relations, such as the negative correlations between BV and T [86, 147, 148], and between I and V [55], also consistently emerge in both models. The main difference between both network structures is the relation between F and A (R4): familiarity only appears to contribute to the arousing quality of positive odors, but not of negative odors. The similarity between the UOEA and POEA networks suggests the existence of multiple affective modes [149]. This notion is reflected in bivariate models of valence (e.g., [150]) that represent pleasant and unpleasant feelings along two separate unipolar dimensions [151153]. Future studies on mixed affective responses to odors may provide more evidence about the dual nature and characteristics of the systems mediating the affective appraisal of odors [154].

We found that the valence component of baseline affective state correlates strongly and negatively with the odor detection threshold (R12). Although this result agrees with the findings that (1) negative mental states raise odor detection thresholds [54, 86] and (2) a positive mental state can lower detection thresholds [86], this relation has (to the best of our knowledge) not been reported previously. Future studies are needed to investigate whether this relation can be replicated with different odors and populations.

In contrast to previous studies that reported no consistent relation between familiarity and valence (R3) for unpleasant odors [25, 33, 51], we find that R3 is positive, independent of odor valence. This discrepancy most likely arises from the fact that these earlier studies computed the correlations at the group level, while we use individual standardizing (i.e., the data for each individual is standardized before computing the networks, to retain a within-individual approach to the data). Thus, it appears that at the individual level, increasing familiarity (i.e., a reduction of uncertainty) consistently enhances valence, both for pleasant odors (valence becomes more positive) and for unpleasant ones (valence becomes less negative). This agrees with the general tendency to attribute more weight to affective information in conditions of uncertainty [155]. Unpleasant odors that are unfamiliar (i.e., for which it has not yet been established whether they are harmful) may be rated as more unpleasant than unpleasant odors that are more familiar and known to be harmless. Our current finding also seems to agree with the finding that the unpleasant odors of fish and garlic were rated as less unpleasant by children who correctly identified them [122].

Contrary to our expectations, we found no negative correlation between T and I (R13).

The high degree of centrality of I in the UOEA network model suggests that I is the most crucial factor influencing the affective appraisal of unpleasant odors. The same holds for F in the POEA network model. T is only affected by BV and appears to have no effect on any of the other variables.

Partial correlation networks are exploratory hypothesis-generating structures that are merely indicative of potential causal effects [104]. With this caveat in mind, the emerging association between BV and T can still be interpreted as a causal relation, due to the temporality of the associated measurements (i.e., the assessment of BV precedes the measurement of T). Insofar as the centrality of a node can be taken to reflect the causal connections emanating from that node, it appears that I dominates the affective appraisal of unpleasant odors (R1, R2), while familiarity dominates the affective appraisal of pleasant odors (R3, R4).

4.1 Limitations

Several limitations should be acknowledged for the present study.

Except for the edges connecting the baseline affective state (BV and BA), the edges in our networks of partial correlation coefficients are undirected and therefore preclude any conclusions about causal (unidirectional or reciprocal) relations. Although correlation does not establish causation, it is consistent with it. Also, the absence of an edge between two variables provides evidence that they are not causally related. Hence, although our exploratory models are in no way confirmatory of causal relationships, they can serve to inspire targeted experimental studies investigating possible predictive relationships. Future research could expand network analysis with a Bayesian network approach [143, 156] to investigate the causal relationships between the different parameters involved in affective odor perception.

In this study we measured olfactory sensitivity using the validated Sniffin’ Sticks based threshold test (SST), which has become a popular (validated and standard) procedure in the literature. This test was also used in several references cited in this study, allowing us to relate our current finding to those earlier results. However, the SST is a single-molecule based general diagnostic tool to assess olfactory functioning. Even for people with a normal sense of smell, sensitivity can vary significantly between individual odors [157]. Hence, the SST measurements cannot be translated to different odors [157] and the test cannot distinguish between general smell dysfunction and PEA insensitivity [158]. This may be the reason why no relation between T and I appears in our results. Future studies could investigate the relationship between threshold and intensity more closely by measuring a detection threshold for each individual odor that is used. However, such a procedure will be very time consuming and tiresome for the participants. Alternatively, future studies may also consider to obtain olfactory threshold assessments that are less dependent on the individual variability in sensitivity to specific odorants by using a threshold test based on complex odor mixtures (e.g. SMELL-R: [158]; see also [159]). However, this invariance only appears to hold for specific odor mixtures [160], and there is currently no generally accepted and validated complex-odor based threshold test available.

The absence of connections between observed variables (nodes) in our networks can either imply that these variables are statically independent when conditioning on all other variables, or it can mean that there was simply insufficient power to detect a relation between these variables [161]. The betweenness centrality estimates were insufficiently stable and should be therefore interpreted cautiously (i.e., the order induced on the nodes by betweenness is not very meaningful). The stability of betweenness centrality might have been greater if we had tested more participants. Future studies including a larger number of participants are required to resolve these issues.

Using Gaussian graphical models, we implicitly assumed that the variables in our models are linearly related. Diagnostic scatterplots show that this assumption is met within the groups of pleasant and unpleasant odors. Although the use of a different kind of correlation estimate (e.g., Spearman of Kendall) would allow for modelling non-linear relations (such as the one shown in Fig 4) these models would be less optimal and new methods are needed to construct a fitting network model.

The current study is a first attempt to construct a network representation for some the main factors that significantly influence the affective appraisal of odors. Given that we mainly focused on a limited set of mediators (intensity, familiarity, sensitivity and the baseline affective state of the observer), we may have missed other relevant factors. For instance, we did not investigate the effects of other factors known to influence the affective appraisal of odors, such as attention [19] or inter-individual variations like differences in physiological state (hunger, satiety: [162]), sex [163], age, semantic knowledge and cultural background [73, 99]. Future studies should investigate how these factors affect network models of odor evoked affect.

In conclusion, these limitations notwithstanding, this study demonstrates that psychometric network analysis can be an effective technique towards the construction of an integral model for the relations between the various factors that influence the affective appraisal of odors. Such a model may constitute the basis for implementing targeted investigations of the way in a wide range of user characteristics determine the affective appraisal of odors.

Supporting information

S1 Data

(XLSX)

Acknowledgments

The authors thank Jos van den Enden (Retroscent, https://retroscent.com) for kindly providing us with a set of additional odorant samples.

They are also grateful for the many comments and suggestions of the anonymous reviewers, that significantly helped to improve the quality of this study.

Data Availability

A CSV file with all results and the R code used to construct the network models are available from the OSF Repository at https://osf.io/psvcy/ with DOI: 10.17605/OSF.IO/PSVCY.

Funding Statement

The authors received no specific funding for this work.

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Decision Letter 0

Alberto Greco

20 Mar 2020

PONE-D-19-24559

A network model of affective odor perception

PLOS ONE

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Reviewer #2: Partly

Reviewer #3: Yes

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2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #2: No

Reviewer #3: I Don't Know

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Reviewer #2: Yes

Reviewer #3: Yes

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5. Review Comments to the Author

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Reviewer #2: The aim of the authors is to systematically investigate the relationships between affective and perceptive judgements of odors, mood and olfactory capacities. Unfortunately and for the reasons I develop below, I do not think that they can claim a substantial contribution for this purpose.

First of all, the tool they use to measure mood is not suitable for fine mood measurement. The authors should better justify the use of EmojiGrid to characterize the mood of participants. It is obvious that positioning one's mood on a two-dimensional space of valence and arousal allows to obtain global information on this mood. But this same space reduces the possibility to dissociate qualitatively different mood states. This is why standardized mood questionnaires such as POMS (Profile of Mood States) exist, which contains more than two valence and arousal scales. Using the EmojiGrid, how to dissociate a state of anxiety (negative valence and high arousal) from an irritable state (negative valence and high arousal). I think you would agree that these moods are not identical and could influence olfactory perception in a specific way. This is problematic because the authors aim to make the link between mood, olfactory perception and olfactory emotional response. How can they answer this question if they do not measure mood according to scales that capture best the quantitative and qualitative differences in mood?

See for instance: “Profile of Mood States.” McNair, D. M., Lorr, M., & Droppleman, L. F. (1992). Revised Manual for the Profile of Mood States. San Diego, CA: Educational and Industrial Testing Services., no. 65 (37),

Second, using the detection threshold of a single molecule in the model is problematic. The authors determine the AEP detection threshold for each participant and use this value to establish the influence of the threshold on odour-related emotional response. This could have been a very important contribution if the authors had performed the detection threshold for each odour used in the affective assessment (I can absolutely not blame the authors for not doing it because it would be an extremely tedious job). Thresholds are specific to each molecule; determining a perception threshold for one molecule does not predict thresholds for others. The test used for the threshold is effective in clinical investigation to detect possible anosmia or hyposmia. This allows the patient to be quickly located according to a known standard. But the approach of the authors in this research is to observe systematic relations between detection thresholds, perceptive and affective dimensions thanks to the use of 40 different odorants. It is not by introducing the threshold to one and only one molecule that these systematic relationships can be studied. One argument for my criticism is that the network analysis the authors performed does not reveal a relationship between threshold and perceived intensity. If the threshold measurement with one molecule was a good estimate of overall olfactory capacity, then a strong positive relationship would have been expected.

Hsieh, J. W., Keller, A., Wong, M., Jiang, R. S., & Vosshall, L. B. (2017). SMELL-S and SMELL-R: olfactory tests not influenced by odor-specific insensitivity or prior olfactory experience. Proceedings of the National Academy of Sciences, 114(43), 11275-11284.

There is, in my opinion, a set of theoretical or definitional clarifications that should be made in the introduction to make the authors' purpose clearer to the reader:

Line 27: Just because there is a high degree of overlap between the cerebral systems that process olfaction and emotions does not mean that the responses have to be "various", this overlap could exist with only one emotional response. Please rephrase or delete that attempted explanation.

Line 30: These dimensions are not necessarily the most fundamental, they are fundamental but not the only ones. They are especially the most used to measure the "core affect". Fontaine, J. R., Scherer, K. R., Roesch, E. B., & Ellsworth, P. C. (2007). The world of emotions is not two-dimensional. Psychological science, 18(12), 1050-1057.

Line 32 and following: I don't understand why the authors use the term "determined" in this sentence, it suggests that there is the valence and arousal of smell (are these intrinsic properties of smell?) that cause the emotional response. Valence and arousal are dimensions of the affective response. The authors may mean that it is possible to describe or characterize the affective response through the psychological dimensions of valence and arousal. There is abundant literature that describes the emotional response as multi-componential, with its subjective, physiological, motivational, cognitive and expressive dimensions. The emotional response cannot be limited to the subjective dimensions of valence and arousal. I suggest that the authors reduce their rationale to the characterization of the relationships between the subjective dimensions of olfactory evaluations.

It seems that the authors differentiate between the intensity of odour, which would be a subjective quality and which they name “perceived intensity”, while they might suggest that the other dimensions represent intrinsic properties of odour, which of course are not. I think it is important to clarify that from the moment that participants are asked to evaluate odour on any dimension, this dimension is subjective and deserves the qualifier "perceived". Thus, the authors could remove the mention of “perceived” in “perceived intensity” or, on the contrary, add this mention to the other dimensions.

Concerning the data availability and analysis strategy.

1. All edges should be undirected except those from Mood which should go from mood to the other variables. Authors can’t postulate mutual relationship in this case because they don’t measure the effect of each odor on participant’s mood.

2. Authors used Gaussian graphical model but didn’t give evidence there data continuous data are distributed. By deliberately selecting a group of positive and a group of negative odours, I fear that the authors have favoured non-normal valence distributions.

3. It seems to me that the measurements are repeated in the same individual (all smells are judged by each individual). Why didn't the authors take this factor into account when applying the GGM? In the model presented by the authors, it seems to me that each observation is considered unique and independent (cross-sectional data analysis) which is not the case.

4. I particularly appreciate the willingness of the authors to share the data from this experiment along with the analysis scripts. Indeed, this is a practice that must now be encouraged if science is to evolve in complete transparency. However, in the data shared by the authors, participant numbers and odours’ names are not reported, which limits the verification and exploitation of the data. I strongly encourage the authors to provide this information which will be very important for the olfactory research community.

Discussion.

The authors conducted the analyses separately for positive and negative odours to “avoid neutralization effects, and because of the functional dichotomy that may exist in the relation between valence and the other variables that are measured in this study (e.g., the relation between valence and familiarity: [37])” The results they present suggest that pleasant and unpleasant networks do not differ. What are the functional consequences of this result? What does this result tell us about the nature of the relationships between the subjective odour dimensions or the treatment of pleasant and unpleasant odours? There are still some small differences between the two networks but it is very difficult for the reader to imagine the importance of these differences.

Reviewer #3: The authors modelled the affective appraisal of odors using sensory and affective parameters using network analysis. This is an interesting approach, especially because current mood is taken into account with references to Clore and Schwarz. Although I feel a model of affective odor appraisal should probably include additional contextual factors going beyond mood; mood can be considered context, and so this is an important step towards an integral model on affective odor appraisal. The authors do point out that future models should include gender and personality, for example. The authors should acknowledge that this may change the relations in the model quite a bit. For example, risk perception from odor exposure or source of exposure will affect the relations.

I have a number of minor to major comments and concerns, that I list below:

Introduction

I am missing a bit of nuance in certain paragraphs like 1.2.1 on perceived intensity. For example, the suggested relation between detection thresholds, intensity and valence should receive more attention. One could interpret the paragraph as implying that individuals with high thresholds show lower intensity perception or different valence perception. In view of the importance of perceived intensity in this paper, please extend this paragraph to include a section on threshold, determinants of perceived intensity and their relation.

There is more attention going to familiarity in 1.2.2, but what I miss is a clear definition of familiarity and how it relates to recognition/identification because it is my impression these terms are mixed up later on in the paper. I would recommend the authors refer to Kösters misfit theory.

1.3 Valence and arousal

What is not addressed here (unless I missed something) is the importance of learning. The general notion in the (human) olfaction literature is that odor valence, with perhaps very few exceptions, is the result of learning, rather than “innate” or “labelled line”. So the context in which an odor was experienced will determine how that odor is liked. While for many individuals context may be similar (e.g. delicious foods) resulting in same direction of valence, the resulting valence may vary (considerably) between individuals. It is important to address this in the paper, to set the stage for the modelling. This is to complete the set of factors we would ideally deal with.

Methods

I am not an expert on network analysis. Since the authors themselves reflect on sample size and (lack of) statistical power in the Discussion, can they reflect on whether the sample size of 56 for this type of analysis? How does this reflect on model fit and related parameters? Have they done a power analysis? I realize testing participants on odor perception in the lab requires substantial effort.

Iso-intensity

Testing for iso-intensity of the stimulus set requires more than three authors sniffing the stimuli to agree on intensity. Typically, this would require preparing multiple concentrations of the stimuli, having panelists smell and rate them and titrate the concentrations to get to iso-intensity over multiple sessions. Can the authors elaborate on what they did to achieve iso-intensity?

However, I am confused as to the need for iso-intensity. Intensity is a factor in the model and therefore it needs to vary. Judging from the outcome, it does, which would contradict the earlier statement of iso-intensity. Why would you need equal intensity in the first place? Perhaps the authors can clarify this.

Table 1: Please include concentrations (% compound in diluent or otherwise) in the Table for replication purposes. What did the safety assessment of compounds like tar, turpentine, and diesel fumes entail?

Procedure (page 14):

- It is always advised to have repetitions on all odor ratings. However, unless I missed this, I guess with 40 odors this was not possible.

Results

- Please provide a Table with means, SD or thresholds and other ratings for all odorants.

- How are individuals who do not judge peach as pleasant, perceive pineapple as lowly intense and peppermint as not familiar, outliers? There is no wrong or right answer as to how pleasant peach should be. See my previous comment about learning. These responses are not “erroneous” (page 20) so please remove. Outlier exclusion is only acceptable if a standard from exclusion is given based on e.g. 2-3 SD’s, so if there is a statistical reason as you model would be radically different perhaps if you included those.

- I am surprised by the finding that mood valence correlates strongly and negatively with threshold. I do not recall having ever seen this relation between lower threshold and positive mood before, even when the authors cite reference 60. I feel it would be important to test the model on another data set of different individuals/odors, so on new data. Will these relations hold? It would have been stronger if separate data sets had been used for model development and testing. But perhaps this is not a common approach for this type of data-driven analysis? Can the authors address this in the Discussion?

Finally, it is “Utrecht University” and not “University of Utrecht” (www.uu.nl – English)

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Decision Letter 1

Alberto Greco

15 Jun 2020

PONE-D-19-24559R1

A network model of affective odor perception

PLOS ONE

Dear Dr. Alexander Toet

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Despite the improvement of the revised paper, there are still some major critical issues to be addressed. Specifically, the authors should better justify their model assumptions and how they can compare their model with the hypothetic model based on the literature. Finally, they have to solve the other minor points raised by the two reviewers.

Please submit your revised manuscript by 30 July 2020 If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

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If you would like to make changes to your financial disclosure, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

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We look forward to receiving your revised manuscript.

Kind regards,

Alberto Greco

Academic Editor

PLOS ONE

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Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #2: (No Response)

Reviewer #3: (No Response)

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2. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #2: Yes

Reviewer #3: Partly

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3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #2: Yes

Reviewer #3: I Don't Know

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4. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #2: Yes

Reviewer #3: Yes

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5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #2: Yes

Reviewer #3: Yes

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6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #2: I think the authors have done an important work to improve the quality of the manuscript. The results presented here make it possible to introduce this family of network analyses to a community of researchers that has not used these techniques much so far. Anyway, I still have a few points I would like to see discussed/modified by the authors before I recommend it for publication.

General major points:

Since the hypothetic model is based on the literature and the model obtained in this study on experimental data related to a sampling of specific odours and participants, how can a meaningful comparison be made? Are the differences not explicable only in terms of sampling? Moreover, the hypothetical model proposed by the authors does not separate pleasant from unpleasant odours (it is crucial for some relations like R3). This makes any comparison very difficult. Why not just removing the theoretical model just report the existing relationships. Or the theoretical model must be separated according to the valence of the odours.

The analyses performed are based on linear correlations between the variables. Don't the authors think that this forced them to exclude any relation of another order (quadratic etc..) and therefore: i) to have to separate pleasant from unpleasant odours and ii) to conclude that there are two different treatments for the different valences. In other words, does the conclusion that pleasant and unpleasant odours must have specific treatments not simply follow from the fact that the analytical techniques employed examine linear relationships? I think the authors should discuss this point (linearity)a little bit more within the limitations.

Specific points:

Line 30 to 36: The three sentences introduce the same notion. Consider rewording them to make the information less redundant.

Line 43 and following: I don't quite understand the authors' inference that "These physiological parameters are linked to arousal..." It is a fairly robust finding in the psychophysiological literature that electrodermal activity is sensitive to arousal but that cardiac variations can be related to valence. An inverted U-curve between the skin conductance responses amplitude and the valence is very often observed for smells but also for images and sounds. This is interpreted as a reflection that subjective arousal and subjective valence are not independent.

Alaoui-Ismaili, O., Robin, O., Rada, H., Dittmar, A., & Vernet-Maury, E. (1997). Basic emotions evoked by odorants: Comparison between autonomic responses and self-evaluation. Physiology and Behavior, 62, 713–720.

Alaoui-Ismaili, O., Vernet-Maury, E., Dittmar, A., Delhomme, G., & Chanel, J. (1997). Odor hedonics: Connection with emotional response estimated by autonomic parameters. Chemical Senses, 22, 237–248.

Bensafi, M., Rouby, C., Farget, V., Bertrand, B., Vigouroux, M., & Holley, A. (2002a). Autonomic nervous system responses to odors: The role of pleasantness and arousal. Chemical Senses, 27, 703–709.

Bensafi, M., Rouby, C., Farget, V., Bertrand, B., Vigouroux, M., & Holley, A. (2002b). Influence of affective and cognitive judgments on autonomic parameters during inhalation of pleasant and unpleasant odors in humans. Neuroscience Letters, 319, 162–166.

Delplanque, S., Grandjean, D., Chrea, C.,Coppin, G.,Aymard, L.,Cayeux, I.,etal. (2009).Sequential unfolding of novelty and pleasantness appraisals of odors: Evidence from facial electromyography and autonomic reactions. Emotion 9, 316–328.doi:10.1037/a0015369

Lang, P. J., Greenwald, M. K., Bradley, M. M., & Hamm, A. O. (1993). Looking at pictures: Affective, facial, visceral, and behavioral reactions. Psychophysiology, 30, 261–273.

Pichon, A., et al. Sensitivity of Physiological Emotional Measures to Odors Depends on the Product and the Pleasantness Ranges Used. In: Frontiers in Psychology, 2015, vol. 6. doi: 10.3389/fpsyg.2015.01821

Line 56 and following: I confess to having some difficulty with the use of the term "influence" in sentence propositions such as "Factors influencing the affective appraisal of odors". This type of sentence implies a causal relationship between factors that would be primary and the affective appraisal of odour that would depend on these factors. I don't think it is possible to postulate this kind of causal relationship, and the experiment conducted by the authors and the partial correlation analyses do not allow us to infer a causal link. At best, the authors can examine the variables that covariate or correlate with the affective appraisal. Is an odour judged more intense then another because it is appraised as more unpleasant? Or is it the contrary? I will come back to this hidden “causal” concept later in my review.

Line 102: The sentence does not give a clear definition of familiarity. I also have difficulty understanding what the authors mean when they write: “Familiarity is implicitly linked to the affective and hedonic appreciation”. Initially, the authors seem to differentiate between affective and hedonic, but there is no mention of differences in the manuscript. Is it necessary to make this distinction? Second, can familiarity be defined without resorting to the hedonic value? Here is a definition that the authors can put in the context of olfaction: “A form of remembering in which a situation, event, place, person, or the like provokes a subjective feeling of recognition and is therefore believed to be in memory, although it is not specifically recalled.”

Line 114: “This suggest that pleasant and unpleasant odors are evaluated by different processing channels, in agreement with models that suggest that different evaluative channels are involved in the processing of negatively and positively valenced stimuli” This sentence doesn't add much with the two parts together. Please consider removing the first part.

Line 279: is it an arithmetic or geometric mean?

Line 388: is the parameter λ the sparsity parameter? Is so please mention it.

Line 419: Please check/correct the definition of betweenness.

Line 536, Table 2: Looking more closely (thanks to the excel file the authors provided) at the correlation between valence and familiarity for unpleasant odours only, conventional correlational analyses give no statistically significant relationship but the correlation coefficient is negative. This result is consistent with several studies in the literature cited by the authors failing to demonstrate any relationship between familiarity and valence for unpleasant odors. How then can we explain that a significant positive relationship appears in network analyses (R3, UPOEA) for the very same odors?

Line 548: typo () -> )

Line 557: For pleasant smells, familiarity and valence differ only in the strength parameter. How do we know if this is different enough to conclude that familiarity governs positive odour ratings? Is the answer at line 612 (“we cannot confidently conclude that any node in the PUOEA network is significantly more central than any other”)?

Line 597 and following: I have some difficulty understanding the authors' reasoning. Do they mean that the sample mean being included in the CI of the bootstrap mean, then the accuracy is good? And then because positive and negative edge so not share CI, they are clearly different? Please reformulate if possible.

Line 691: please add a nuance like “as measured in term of valence and arousal” because there are other emotion theories (not only dimensional models) and you could have chosen to assess affective appraisal via rating on basic emotions.

Line 718: The authors should not link the networks they highlight at the level of the subjective variables measured to any brain functioning. The structure of the networks they obtain can in no way reflect any brain functioning. Their conclusions should be limited to associative networks only.

Line 744 and following: I thought partial correlations can be indicative of potential causal pathways. As mentioned by Epskamp and Fried, 2018): “…partial correlation networks are thought of as highly exploratory hypothesis-generating structures, indicative of potential causal effects.” I think that the authors have to take precautions with the notion of causality. Even if it is very clear to them, they should be very careful not to imply that causal links are demonstrated by these analytical techniques. It is only suggested, assuming that there is no hidden latent unmeasured variable. Although the authors discuss this limitation later in the manuscript (in the limitation section). I think that they should not let people think that the causal link is there.

Epskamp, S., & Fried, E. I. (2018). A tutorial on regularized partial correlation networks. Psychological methods, 23(4), 617.

Reviewer #3: Intensities

The authors replied that the purpose of the pilot experiment was not to achieve iso-intensity.

I would agree that this would have been problematic in light of the objective of modelling. For the current objective, it is ideal for the odorants to cover a range of intensities so that there is variation. The authors state that extreme differences in intensity are undesirable and seem to aim for intensities to comparable (page 10). This leaves me in the dark as to the range of intensities they would deem acceptable or comparable. It now appears as if they “just went” with whatever the concentration was that the supplier sent them. Usually, dilutions would be made from those (in general high-intense) odorants the supplier will have prepared, to finetune the range of concentrations to cover a pre-agreed range of intensities.

I understand that with Sniffin’ Sticks already prepared by Burghart (with the odorant already in the Stick) it defies the purpose to have to go back to ordering the chemical and prepare the stick all over again, although these were prepared to be comparable and not vary too much. For odorants ordered from suppliers dilutions to achieve variation could have been made.

The authors have addressed the variation for valence, which clearly differed. Can the authors now address address the variation in intensity and if not by providing data from their pilot, by reflecting on this issue?

For an example of how concentrations determination is done to match intensities please see Keller, A., Hempstead, M., Gomez, I.A. et al. An olfactory demography of a diverse metropolitan population. BMC Neurosci 13, 122 (2012). https://doi.org/10.1186/1471-2202-13-122

for determination of “high” and “low” levels.

I would agree would be exaggerated to put this much work into determination of range for the present paper, but compared with this example I think the authors can do more than they currently do to address this. The means vary between 43 and 77 which strikes me as a bit narrow, but you can convince me otherwise.

Why not insert the table of means in the main paper?

Safety

When it comes to safety levels of tar and diesel fume, the authors now refer the reader to find this out for themselves on the supplier website. It is not the responsibility of the reader to verify safety, but that of the scientists. Based on the supplier information can the authors include a declaration in the paper that levels presented were safe? This is not to be dogmatic, but also to state that including such a statement to show that due diligence has been performed in protecting the participant’s health.

Finally:

I searched through the manuscript to find the difference baseline affective state and current affective state, but now conclude they are the same. If that is indeed the case, it would help to consistently use the same term.

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Reviewer #2: No

Reviewer #3: No

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Decision Letter 2

Alberto Greco

9 Jul 2020

A network model of affective odor perception

PONE-D-19-24559R2

Dear Dr. Alexander Toet,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

An invoice for payment will follow shortly after the formal acceptance. To ensure an efficient process, please log into Editorial Manager at http://www.editorialmanager.com/pone/, click the 'Update My Information' link at the top of the page, and double check that your user information is up-to-date. If you have any billing related questions, please contact our Author Billing department directly at authorbilling@plos.org.

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Kind regards,

Alberto Greco

Academic Editor

PLOS ONE

Additional Editor Comments (optional):

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #2: All comments have been addressed

**********

2. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #2: Yes

**********

3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #2: Yes

**********

4. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #2: Yes

**********

5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #2: Yes

**********

6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #2: I think the authors have correctly addressed the various points raised during the article review process. It is, in my opinion, now acceptable for publication in PLOS ONE. However, I do have one final minor suggestion that may help in understanding the non-trivial concepts of "modules" vs. "modes".

Line 676: The difference between processes by modules or modes is not necessarily obvious to readers. The authors could add a small sentence that defines more clearly and explicitly what differentiates the two.

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7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #2: Yes: Sylvain Delplanque

Acceptance letter

Alberto Greco

15 Jul 2020

PONE-D-19-24559R2

A network model of affective odor perception

Dear Dr. Toet:

I'm pleased to inform you that your manuscript has been deemed suitable for publication in PLOS ONE. Congratulations! Your manuscript is now with our production department.

If your institution or institutions have a press office, please let them know about your upcoming paper now to help maximize its impact. If they'll be preparing press materials, please inform our press team within the next 48 hours. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information please contact onepress@plos.org.

If we can help with anything else, please email us at plosone@plos.org.

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Kind regards,

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on behalf of

Dr. Alberto Greco

Academic Editor

PLOS ONE

Associated Data

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    Data Availability Statement

    A CSV file with all results and the R code used to construct the network models are available from the OSF Repository at https://osf.io/psvcy/ with DOI: 10.17605/OSF.IO/PSVCY.


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