Testing detectability, attractivity, hedonic specificity, extractability, and robustness of colostrum odor-Toward an olfactory bioassay for human neonates.

Human milk odor is attractive and appetitive for human newborns. Here, we studied behavioral and heart-rate (HR) responses of 2-day-old neonates to the odor of human colostrum. To evaluate detection in two conditions of stimulus delivery, we first presented the odor of total colostrum against water. Second, the hedonic specificity of total colostrum odor was tested against vanilla odor. Third, we delivered only the fresh effluvium of colostrum separated from the colostrum matrix; the stability of this colostrum effluvium was then tested after deep congelation; finally, after sorptive extraction of fresh colostrum headspace, we assessed the activity of colostrum volatiles eluting from the gas chromatograph (GC). Regardless of the stimulus-delivery method, neonates displayed attraction reactions (HR decrease) as well as appetitive oral responses to the odor of total colostrum but not to vanilla odor. The effluvium separated from the fresh colostrum matrix remained appetitive but appeared labile under deep freezing. Finally, volatiles from fresh colostrum effluvium remained behaviorally active after GC elution, although at lower magnitude. In sum, fresh colostrum effluvium and its eluate elicited a consistent increase in newborns' oral activity (relative to water or vanilla), and they induced shallow HR decrease. Newborns' appetitive oral behavior was the most reproducible response criterion to the effluvium of colostrum. In conclusion, a set of unidentified volatile compounds from human colostrum is robust enough after extraction from the original matrix and chromatographic processing to continue eliciting appetitive responses in neonates, thus opening new directions to isolate and assay specific volatile molecules of colostrum.

The spatial distribution of eye movements predicts the (false) recognition of emotional facial expressions.

Recognizing facial expressions of emotions is a fundamental ability for adaptation to the social environment. To date, it remains unclear whether the spatial distribution of eye movements predicts accurate recognition or, on the contrary, confusion in the recognition of facial emotions. In the present study, we asked participants to recognize facial emotions while monitoring their gaze behavior using eye-tracking technology. In Experiment 1a, 40 participants (20 women) performed a classic facial emotion recognition task with a 5-choice procedure (anger, disgust, fear, happiness, sadness). In Experiment 1b, a second group of 40 participants (20 women) was exposed to the same materials and procedure except that they were instructed to say whether (i.e., Yes/No response) the face expressed a specific emotion (e.g., anger), with the five emotion categories tested in distinct blocks. In Experiment 2, two groups of 32 participants performed the same task as in Experiment 1a while exposed to partial facial expressions composed of actions units (AUs) present or absent in some parts of the face (top, middle, or bottom). The coding of the AUs produced by the models showed complex facial configurations for most emotional expressions, with several AUs in common. Eye-tracking data indicated that relevant facial actions were actively gazed at by the decoders during both accurate recognition and errors. False recognition was mainly associated with the additional visual exploration of less relevant facial actions in regions containing ambiguous AUs or AUs relevant to other emotional expressions. Finally, the recognition of facial emotions from partial expressions showed that no single facial actions were necessary to effectively communicate an emotional state. In contrast, the recognition of facial emotions relied on the integration of a complex set of facial cues.

Human neonates prefer colostrum to mature milk: Evidence for an olfactory bias toward the "initial milk"?

OBJECTIVES: Colostrum is the initial milk secretion which ingestion by neonates warrants their adaptive start in life. Colostrum is accordingly expected to be attractive to newborns. The present study aims to assess whether colostrum is olfactorily attractive for 2-day-old newborns when presented against mature milk or a control. METHODS: The head-orientation of waking newborns was videotaped in three experiments pairing the odors of: (a) colostrum (sampled on postpartum day 2, not from own mother) and mature milk (sampled on average on postpartum day 32, not from own mother) (n tested newborns = 15); (b) Colostrum and control (water; n = 9); and (c) Mature milk and control (n = 13). RESULTS: When facing the odors of colostrum and mature milk, the infants turned their nose significantly longer toward former (32.8 vs 17.7% of a 120-s test). When exposed to colostrum against the control, they responded in favor of colostrum (32.9 vs 16.6%). Finally, when the odor of mature milk was presented against the control, their response appeared undifferentiated (26.7 vs 28.6%). CONCLUSIONS: These results indicate that human newborns can olfactorily differentiate conspecific lacteal fluids sampled at different lactation stages. They prefer the odor of the mammary secretion - colostrum - collected at the lactation stage that best matches the postpartum age of their own mother. These results are discussed in the context of the earliest mother-infant chemo-communication. Coinciding maternal emission and offspring reception of chemosignals conveyed in colostrum may be part of the sensory precursors of attunement between mothers and infants.

Olfaction scaffolds the developing human from neonate to adolescent and beyond.

The impact of the olfactory sense is regularly apparent across development. The fetus is bathed in amniotic fluid (AF) that conveys the mother's chemical ecology. Transnatal olfactory continuity between the odours of AF and milk assists in the transition to nursing. At the same time, odours emanating from the mammary areas provoke appetitive responses in newborns. Odours experienced from the mother's diet during breastfeeding, and from practices such as pre-mastication, may assist in the dietary transition at weaning. In parallel, infants are attracted to and recognize their mother's odours; later, children are able to recognize other kin and peers based on their odours. Familiar odours, such as those of the mother, regulate the child's emotions, and scaffold perception and learning through non-olfactory senses. During juvenility and adolescence, individuals become more sensitive to some bodily odours, while the timing of adolescence itself has been speculated to draw from the chemical ecology of the family unit. Odours learnt early in life and within the family niche continue to influence preferences as mate choice becomes relevant. Olfaction thus appears significant in turning on, sustaining and, in cases when mother odour is altered, disturbing adaptive reciprocity between offspring and carer during the multiple transitions of development between birth and adolescence. This article is part of the Theo Murphy meeting issue 'Olfactory communication in humans'.

Dysfunction of the Mesolimbic Circuit to Food Odors in Women With Anorexia and Bulimia Nervosa: A fMRI Study.

Brain reward dysfunction in eating disorders has been widely reported. However, whether the neural correlates of hedonic and motivational experiences related to food cues are differentially affected in anorexia nervosa of restrictive type (ANr), bulimia nervosa (BN), and healthy control (HC) participants remains unknown. Here, 39 women (14 ANr, 13 BN, and 12 HC) underwent fMRI while smelling food or non-food odors in hunger and satiety states during liking and wanting tasks. ANr and BN patients reported less desire to eat odor-cued food and odor-cued high energy-density food (EDF), respectively. ANr patients exhibited lower ventral tegmental area (VTA) activation than BN patients to food odors when rating their desire to eat, suggesting altered incentive salience attribution to food odors. Compared with HC participants, BN patients exhibited decreased activation of the caudate nucleus to food odors in the hunger state during the wanting task. Both patient groups also showed reduced activation of the anterior ventral pallidum and insula in response to high EDF odors in the hunger state during the wanting task. These findings indicate that brain activation within the food reward-regulating circuit differentiates the three groups. ANr patients further exhibited lower activation of the precuneus than other participants, suggesting a possible role of body image distortion in ANr. Our study highlights that food odors are relevant sensory probes to gain better insight into the dysfunction of the mesolimbic and striatal circuitry involved in food reward processing in patients with EDs.

Watching happy faces potentiates incentive salience but not hedonic reactions to palatable food cues in overweight/obese adults.

'Wanting' and 'liking' are mediated by distinct brain reward systems but their dissociation in human appetite and overeating remains debated. Further, the influence of socioemotional cues on food reward is little explored. We examined these issues in overweight/obese (OW/OB) and normal-weight (NW) participants who watched food images varying in palatability in the same time as videoclips of avatars looking at the food images while displaying facial expressions (happy, disgust or neutral) with their gaze directed only toward the food or consecutively toward the food and participants. We measured heart rate (HR) deceleration as an index of attentional/incentive salience, facial EMG activity as an index of hedonic or disgust reactions, and self-report of wanting and liking. OW/OB participants exhibited a larger HR deceleration to palatable food pictures than NW participants suggesting that they attributed greater incentive salience to food cues. However, in contrast to NW participants, they did not display increased hedonic facial reactions to the liked food cues. Subjective ratings of wanting and liking did not differentiate the two groups. Further, OW/OB participants had more pronounced HR deceleration than NW participants to palatable food cues when they watched avatars' happy faces gazing at the food. In line with the "incentive-sensitization" hypothesis, our data suggest that incentive salience attribution and not hedonic reactivity is increased in OW/OB individuals and that happy faces, as social reward cues, potentiate implicit wanting in OW/OB people.

Mimicking emotions: how 3-12-month-old infants use the facial expressions and eyes of a model.

While there is an extensive literature on the tendency to mimic emotional expressions in adults, it is unclear how this skill emerges and develops over time. Specifically, it is unclear whether infants mimic discrete emotion-related facial actions, whether their facial displays are moderated by contextual cues and whether infants' emotional mimicry is constrained by developmental changes in the ability to discriminate emotions. We therefore investigate these questions using Baby-FACS to code infants' facial displays and eye-movement tracking to examine infants' looking times at facial expressions. Three-, 7-, and 12-month-old participants were exposed to dynamic facial expressions (joy, anger, fear, disgust, sadness) of a virtual model which either looked at the infant or had an averted gaze. Infants did not match emotion-specific facial actions shown by the model, but they produced valence-congruent facial responses to the distinct expressions. Furthermore, only the 7- and 12-month-olds displayed negative responses to the model's negative expressions and they looked more at areas of the face recruiting facial actions involved in specific expressions. Our results suggest that valence-congruent expressions emerge in infancy during a period where the decoding of facial expressions becomes increasingly sensitive to the social signal value of emotions.

Responses of Human Neonates to Highly Diluted Odorants from Sweat.

Conjugated forms of odorants contributing to sweat odor occur not only in human sweat but also in amniotic fluid, colostrum, and milk. However, it is unclear whether the released odorants are detected and hedonically discriminated by human newborns. To investigate this issue, we administered highly diluted solutions of (R)/(S)-3-methyl-3-sulfanylhexan-1-ol (MSH), (R)/(S)-3-sulfanylhexan-1-ol (SH), (E)/(Z)-3-methylhex-2-enoic acid (3M2H), and (R)/(S)-3-hydroxy-3-methylhexanoic acid (HMHA) to 3-d-old infants while their respiratory rate and oro-facial movements were recorded. Adult sensitivity to these odorants was assessed via triangle tests. Whereas no neonatal stimulus-specific response was found for respiratory rate, oro-facial reactivity indicated orthonasal detection of MSH and SH by male neonates, and of HMHA by the whole group of neonates. Dependent on the dilution of odorants, newborns evinced neutral responses or longer negative oro-facial expressions compared with the reference stimuli. Finally, newborns appeared to be more sensitive to the target odorants than did adults.

Emotional expressiveness of 5-6 month-old infants born very premature versus full-term at initial exposure to weaning foods.

Facial expressions of 5-6 month-old infants born preterm and at term were compared while tasting for the first time solid foods (two fruit and two vegetable purées) given by the mother. Videotapes of facial reactions to these foods were objectively coded during the first six successive spoons of each test food using Baby FACS and subjectively rated by naïve judges. Infant temperament was also assessed by the parents using the Infant Behaviour Questionnaire. Contrary to our expectations, infants born preterm expressed fewer negative emotions than infants born full-term. Naïve judges rated infants born preterm as displaying more liking than their full-term counterparts when tasting the novel foods. The analysis of facial expressions during the six spoonfuls of four successive meals (at 1-week intervals) suggested a familiarization effect with the frequency of negative expressions decreasing after tasting the second spoon, regardless of infant age, type of food and order of presentation. Finally, positive and negative dimensions of temperament reported by the parents were related with objective and subjective coding of affective reactions toward foods in infants born preterm or full-term. Our research indicates that premature infants are more accepting of novel foods than term infants and this could be used for supporting the development of healthy eating patterns in premature infants. Further research is needed to clarify whether reduced negativity by infants born prematurely to the exposure to novel solid foods reflects a reduction of an adaptive avoidant behaviour during the introduction of novel foods.

Emotional communication in the context of joint attention for food stimuli: effects on attentional and affective processing.

Guided by distinct theoretical frameworks (the embodiment theories, shared-signal hypothesis, and appraisal theories), we examined the effects of gaze direction and emotional expressions (joy, disgust, and neutral) of virtual characters on attention orienting and affective reactivity of participants while they were engaged in joint attention for food stimuli contrasted by preference (disliked, moderately liked, and liked). The participants were exposed to videos of avatars looking at food and displaying facial expressions with their gaze directed either toward the food only or toward the food and participants consecutively. We recorded eye-tracking responses, heart rate, facial electromyography (zygomatic, corrugator, and levator labii regions), and food wanting/liking. The avatars' joy faces increased the participants' zygomatic reactions and food liking, with mutual eye contact boosting attentional responses. Eye contact also fostered disgust reactions to disliked food, regardless of the avatars' expressions. The findings show that joint attention for food accompanied by face-to-face emotional communication elicits differential attentional and affective responses. The findings appear consistent with the appraisal theories of emotion.

Reward for food odors: an fMRI study of liking and wanting as a function of metabolic state and BMI.

Brain reward systems mediate liking and wanting for food reward. Here, we explore the differential involvement of the following structures for these two components: the ventral and dorsal striatopallidal area, orbitofrontal cortex (OFC), anterior insula and anterior cingulate. Twelve healthy female participants were asked to rate pleasantness (liking of food and non-food odors) and the desire to eat (wanting of odor-evoked food) during event-related functional magnetic resonance imaging (fMRI). The subjective ratings and fMRI were performed in hunger and satiety states. Activations of regions of interest were compared as a function of task (liking vs wanting), odor category (food vs non-food) and metabolic state (hunger vs satiety). We found that the nucleus accumbens and ventral pallidum were differentially involved in liking or wanting during the hunger state, which suggests a reciprocal inhibitory influence between these structures. Neural activation of OFC subregions was correlated with either liking or wanting ratings, suggesting an OFC role in reward processing magnitude. Finally, during the hunger state, participants with a high body mass index exhibited less activation in neural structures underlying food reward processing. Our results suggest that food liking and wanting are two separable psychological constructs and may be functionally segregated within the cortico-striatopallidal circuit.

Responsiveness of human neonates to the odor of 5α-androst-16-en-3-one: a behavioral paradox?

The odorous steroid 5α-androst-16-en-3-one (AND) occurs in numerous biological fluids in mammals, including man, where it is believed to play a chemocommunicative role. As AND was recently detected in milk and amniotic fluid, sensitivity and hedonic responses to this substance were assessed in human neonates. To this aim, respiration and facial expressions were recorded in 3-day-old newborns in response to aqueous solutions of AND, ranging from 500ng/mL to 0.5 fg/mL. Although analyses of respiratory rate did not lead to clear-cut results, the newborns changed their facial expressions at concentrations not detected by adults in a triangle test. Newborns displayed negative facial actions of longer duration to AND relative to an odorless control. Thus, AND may be considered to be offensive to newborns, which is a counterintuitive outcome as they are exposed to this compound in the womb (and it should therefore be familiar), in milk, and on the mother's skin surface (and it should therefore be conditioned as positive). Multiple reasons for this perceptual-behavioral paradox are discussed.

Revisiting mu suppression in autism spectrum disorder.

Two aspects of the EEG literature lead us to revisit mu suppression in Autism Spectrum Disorder (ASD). First and despite the fact that the mu rhythm can be functionally segregated in two discrete sub-bands, 8-10 Hz and 10-12/13 Hz, mu-suppression in ASD has been analyzed as a homogeneous phenomenon covering the 8-13 Hz frequency. Second and although alpha-like activity is usually found across the entire scalp, ASD studies of action observation have focused on the central electrodes (C3/C4). The present study was aimed at testing on the whole brain the hypothesis of a functional dissociation of mu and alpha responses to the observation of human actions in ASD according to bandwidths. Electroencephalographic (EEG) mu and alpha responses to execution and observation of hand gestures were recorded on the whole scalp in high functioning subjects with ASD and typical subjects. When two bandwidths of the alpha-mu 8-13 Hz were distinguished, a different mu response to observation appeared for subjects with ASD in the upper sub-band over the sensorimotor cortex, whilst the lower sub-band responded similarly in the two groups. Source reconstructions demonstrated that this effect was related to a joint mu-suppression deficit over the occipito-parietal regions and an increase over the frontal regions. These findings suggest peculiarities in top-down response modulation in ASD and question the claim of a global dysfunction of the MNS in autism. This research also advocates for the use of finer grained analyses at both spatial and spectral levels for future directions in neurophysiological accounts of autism.

Evidence for unintentional emotional contagion beyond dyads.

Little is known about the spread of emotions beyond dyads. Yet, it is of importance for explaining the emergence of crowd behaviors. Here, we experimentally addressed whether emotional homogeneity within a crowd might result from a cascade of local emotional transmissions where the perception of another's emotional expression produces, in the observer's face and body, sufficient information to allow for the transmission of the emotion to a third party. We reproduced a minimal element of a crowd situation and recorded the facial electromyographic activity and the skin conductance response of an individual C observing the face of an individual B watching an individual A displaying either joy or fear full body expressions. Critically, individual B did not know that she was being watched. We show that emotions of joy and fear displayed by A were spontaneously transmitted to C through B, even when the emotional information available in B's faces could not be explicitly recognized. These findings demonstrate that one is tuned to react to others' emotional signals and to unintentionally produce subtle but sufficient emotional cues to induce emotional states in others. This phenomenon could be the mark of a spontaneous cooperative behavior whose function is to communicate survival-value information to conspecifics.

Children's reward responses to picture- and odor-cued food stimuli: a developmental analysis between 6 and 11years.

The reward system is largely involved in the control of food intake. Whether components of this system (i.e., wanting and liking) change during development remains understudied, as well as how proximate factors (sensory cues, motivational state) modulate reward reactivity across development. We examined the developmental pattern of wanting and liking for sensorily-cued food stimuli in 6-11year old children as a function of the child's motivational state (hunger/satiety), gender, and the nature of foods. School children were exposed before or after their lunch on alternative days to visual and odor stimuli representing different categories of familiar foods. Their task was to rate wanting and liking of pictures and odorants of pizza, meat, vegetables, fruits, and chocolate. The following results were found: (1) While liking appeared to be stable from age 6 to 11, more particularly for visually-cued foods, wanting decreased, as well as did subjective hunger perception; (2) there were smaller or absent state-effects in 7-to-9-year-olds; (3) reward ratings were higher in boys than in girls; (4) reward ratings of vegetables were the lowest at all ages. These results suggest that wanting, but not liking, is developmentally variable over childhood, and that this variation depends on age, gender, motivational state (hunger/satiety), the nature of the food and the modality of the sensory cue representing it. Such developmental changes are discussed in relation to biological (adiposity rebound) and cognitive (dietary restraint) factors influencing the motivation to eat during middle (6-7years) and late (9-11years) childhood.

Self-relevance appraisal influences facial reactions to emotional body expressions.

People display facial reactions when exposed to others' emotional expressions, but exactly what mechanism mediates these facial reactions remains a debated issue. In this study, we manipulated two critical perceptual features that contribute to determining the significance of others' emotional expressions: the direction of attention (toward or away from the observer) and the intensity of the emotional display. Electromyographic activity over the corrugator muscle was recorded while participants observed videos of neutral to angry body expressions. Self-directed bodies induced greater corrugator activity than other-directed bodies; additionally corrugator activity was only influenced by the intensity of anger expresssed by self-directed bodies. These data support the hypothesis that rapid facial reactions are the outcome of self-relevant emotional processing.

Self-relevance appraisal of gaze direction and dynamic facial expressions: effects on facial electromyographic and autonomic reactions.

What processes or mechanisms mediate interpersonal matching of facial expressions remains a debated issue. As theoretical approaches to underlying processes (i.e., automatic motor mimicry, communicative intent, and emotional appraisal) make different predictions about whether facial responses to others' facial expressions are influenced by perceived gaze behavior, we examined the impact of gaze direction and dynamic facial expressions on observers' autonomic and rapid facial reactions (RFRs). We recorded facial electromyography activity over 4 muscle regions (Corrugator Supercilli, Zygomaticus Major, Lateral Frontalis, and Depressor Anguli Oris), skin conductance response and heart rate changes in participants passively exposed to virtual characters displaying approach-oriented (anger and happiness), and avoidance-oriented (fear and sadness) emotion expressions with gaze either directed at or averted from the observer. Consistent with appraisal theories, RFRs were potentiated by mutual eye contact when participants viewed happy and angry expressions, while RFRs occurred only to fear expressions with averted gaze. RFRs to sad expressions were not affected by gaze direction. The interaction between emotional expressions and gaze direction was moderated by participants' gender. The pattern of autonomic reactivity was consistent with the view that salient social stimuli increase physiological arousal and attentional resources, with gaze direction, nature of emotion, and gender having moderating effects. These results suggest the critical role of self-relevance appraisal of senders' contextual perceptual cues and individual characteristics to account for interpersonal matching of facial displays.

Does the brain know who is at the origin of what in an imitative interaction?

Brain correlates of the sense of agency have recently received increased attention. However, the explorations remain largely restricted to the study of brains in isolation. The prototypical paradigm used so far consists of manipulating visual perception of own action while asking the subject to draw a distinction between self- versus externally caused action. However, the recent definition of agency as a multifactorial phenomenon combining bottom-up and top-down processes suggests the exploration of more complex situations. Notably there is a need of accounting for the dynamics of agency in a two-body context where we often experience the double faceted question of who is at the origin of what in an ongoing interaction. In a dyadic context of role switching indeed, each partner can feel body ownership, share a sense of agency and altogether alternate an ascription of the primacy of action to self and to other. To explore the brain correlates of these different aspects of agency, we recorded with dual EEG and video set-ups 22 subjects interacting via spontaneous versus induced imitation (II) of hand movements. The differences between the two conditions lie in the fact that the roles are either externally attributed (induced condition) or result from a negotiation between subjects (spontaneous condition). Results demonstrate dissociations between self- and other-ascription of action primacy in delta, alpha and beta frequency bands during the condition of II. By contrast a similar increase in the low gamma frequency band (38-47 Hz) was observed over the centro-parietal regions for the two roles in spontaneous imitation (SI). Taken together, the results highlight the different brain correlates of agency at play during live interactions.

Orofacial reactivity to the sight and smell of food stimuli. Evidence for anticipatory liking related to food reward cues in overweight children.

Whether food liking may be a risk factor of overconsumption and overweight/obesity remains a controversial issue. So far, most studies used subjective reports to assess consummatory behavior, approaches that might overlook subtle or implicit hedonic changes to sensory properties of foods. Therefore, we used a cue-exposure approach by recording different measures of hedonic processes (orofacial reactivity, self-rated pleasantness, food preference) in 6-11 years old overweight (n=20) and normal-weight (n=20) children. Children were exposed to the smell and sight of high and low-energy density food stimuli and to non-food stimuli during pre- and post-prandial states. Their facial and verbal responses were videotaped and parent's reports of children's eating styles and appetitive traits were collected using the Children's Eating Behavior Questionnaire (CEBQ). Results showed that orofacial reactivity, as an objective measure of anticipatory liking, was more discriminative than self reports, with overweight children displaying more lip sucking than normal-weight children when exposed to high energy food pictures and to food odorants. Orofacial reactivity to food cues was also associated with BMI and children's eating styles (food responsiveness, emotional overeating, and desire to drink). Finally, overweight children classified more frequently non-food odorants as members of the food category during the pre-prandial state than during the post-prandial state, suggesting a possible influence of affective/motivational bias on odor categorization. Our findings suggest that orofacial responsiveness may be relevant to assess the sensitivity to energy-dense food reward cues in overweight children and for signaling, as an index of anticipatory liking, a potential risk for the development of overweight/obesity.

An overlooked aspect of the human breast: areolar glands in relation with breastfeeding pattern, neonatal weight gain, and the dynamics of lactation.

The early nursing-sucking relationship is not to be taken for granted in humans. A number of factors can either facilitate or mitigate its optimal establishment on the mother's or newborn's sides. Among these factors, a morphological feature of human mothers' breasts--the areolar glands (AG)--has been identified as potentially important. Three day-old infants display attraction during the presentation of the native secretions of the AG, suggesting that they could influence the newborn's behaviour during breastfeeding. The present study assessed this topic in a sample of 121 Caucasian mother-infant dyads. The areolae of these women were screened during the first 3 postnatal days in parallel with the infant's sucking performance, body weight fluctuations and time to lactation onset. On average, 97% of the women bore AG, 80.2% having 1-20 units per areola and 33% showing AG excreting a visible fluid. The endowment in AG appeared positively linked with neonatal growth after birth and with the speed of lactation onset: infants of primiparous women with lower AG numbers had a lower weight gain than those of mothers with higher AG numbers. Further, it took longer to primiparae with lower AG counts to set on lactation. This study confirms and extends the fact that AG, in interaction with maternal experience, might influence the initiation of the breastfeeding relationship.