Olfactory conditioning by association with histidine-free or balanced amino acid loads in rats

An histidine-devoid but otherwise balanced amino acid mixture depressed food intake from 2 hr after its gastric intubation. It induced conditioned aversion to an odour incorporated in a protein-free diet presented for 6 h following intubation. In other rats, a balanced amino acid mixture established conditioned preference for odour presented in the same diet for 6 h following intubation. The degree of preference was considerably less than the degree of aversion.     

Rat pups prefer their siblings to their mothers: Possible implications for the development of kin recognition

Previous experiments have indicated that for rat pups some learning of their kin's characteristics is required for kin recognition. By differentially scenting the mother and litter with artificial odours, the possible role of each in the development of this ability can be assessed. 12- and 20-day-old rat pups were given a preference tests between the odour associated with their mother and an odour associated with their littermates. It was hypothesized that, because of the importance of the mother to her pups, individuals would preferentially acquire information from her and thus exhibit a preference for that odour. The results, however, did not support this: Both 12- and 20-day-old rat pups were found to prefer their littermate's odour to that of their mother. A further experiment revealed this did not reflect an aversion to their mother's odour. This preference for sibling littermates probably reflects the greater exposure of littermates to one another. The possible advantages of learning about one's kin from littermates are discussed.     

The acquisition of odour qualities

Odours can evoke a large range of qualities. Some of these qualities (e.g., sweetness) appear to be acquired through simultaneously experiencing odours mixed with tastes. Acquisition might also occur when two odours are experienced as a mixture. In this case mixture components might acquire each other's qualities. This was tested in two experiments. In the first, subjects repeatedly sniffed two odour mixtures (either AX, BY or AY, BX). Odours mixed with A acquired A's properties and were judged more similar to A than to B. Odours mixed with B were not clearly discriminable. The second experiment used a similar approach except that Odour B was replaced. Subjects now smelled either AX, CY or AY, CX. All components were discriminable. Odours mixed with A acquired A's properties and were judged more similar to A than to C. Although odours mixed with C did not acquire C's qualities due to a confusion of quality terms, odours previously mixed with C were judged as more similar to C than to A. Evidence of other quality exchanges were also obtained. These results suggest that pure odour qualities can be learnt and lend support to William James's (1890) notion of the acquired equivalence/distinctiveness of cues.     

Experience dependent changes in odour-viscosity perception

One consequence of experiencing flavour — the combination of taste, smell and somatosensation that occurs during ingestion — is that it can result in perceptual changes for the odour component, when this is later smelled alone. One such change is the acquisition of taste-like properties, but whether odours can also acquire somatosensory-like qualities is largely unknown. Participants here were exposed to one odour sampled in a viscous solution, another sampled in a sweet/viscous solution, and a further odour sampled in water. The odour sampled in the sweet/viscous solution was, when later sniffed alone, judged to smell thicker and sweeter, than the other two odours. Similarly, when the sweet/viscous paired odour was added to a viscous solution, the combination was judged as more viscous, than the other two odours — and sweeter when added to a sweet solution. This experiment suggests that odours can acquire tactile-like somatosensory qualities and this may best occur when a taste is present during learning. Recent work indicates that tastes may be superior to somatosensory stimuli alone in promoting flavour binding, a seeming precondition for this type of learning.     

Embryonic olfactory learning in frogs.

We examined the effect of embryonic olfactory experiences on behaviour after hatching in two species of anuran amphibians, Rana temporaria and Rana sylvatica. Odorants (orange and citral) injected into the egg were "learned" by the embryo and subsequently preferred by tadpoles after hatching. The observed preferences were specific to the odours experienced prior to hatching, and tadpoles discriminated between the odour experienced as embryos and a novel odour. The acquired preference was maintained after metamorphosis for those individuals that exhibited a preference as tadpoles. Preferences of tadpoles were also influenced "naturally" by odours present in the water surrounding the developing embryos. The experimentally induced olfactory preferences did not appear to influence the exhibition of sibling recognition abilities. The study represents the first demonstration of "embryonic" learning in amphibians, and the functions and importance of early olfactory learning for amphibians are discussed.     

Peak shift in honey bee olfactory learning

If animals are trained with two similar stimuli such that one is rewarding (S+) and one punishing (S?), then following training animals show a greatest preference not for the S+, but for a novel stimulus that is slightly more different from the S? than the S+ is. This peak shift phenomenon has been widely reported for vertebrates and has recently been demonstrated for bumblebees and honey bees. To explore the nature of peak shift in invertebrates further, here we examined the properties of peak shift in honey bees trained in a free-flight olfactory learning assay. Hexanal and heptanol were mixed in different ratios to create a continuum of odour stimuli. Bees were trained to artificial flowers such that one odour mixture was rewarded with 2 molar sucrose (S+), and one punished with distasteful quinine (S?). After training, bees were given a non-rewarded preference test with five different mixtures of hexanal and heptanol. Following training bees’ maximal preference was for an odour mixture slightly more distinct from the S? than the trained S+. This effect was not seen if bees were initially trained with two distinct odours, replicating the classic features of peak shift reported for vertebrates. We propose a conceptual model of how peak shift might occur in honey bees. We argue that peak shift does not require any higher level of processing than the known olfactory learning circuitry of the bee brain and suggest that peak shift is a very general feature of discrimination learning.     

Embryonic olfactory learning in frogs

We examined the effect of embryonic olfactory experiences on behaviour after hatching in two species of anuran amphibians, Rana temporaria and Rana sylvatica. Odorants (orange and citral) injected into the egg were “learned” by the embryo and subsequently preferred by tadpoles after hatching. The observed preferences were specific to the odours experienced prior to hatching, and tadpoles discriminated between the odour experienced as embryos and a novel odour. The acquired preference was maintained after metamorphosis for those individuals that exhibited a preference as tadpoles. Preferences of tadpoles were also influenced “naturally” by odours present in the water surrounding the developing embryos. The experimentally induced olfactory preferences did not appear to influence the exhibition of sibling recognition abilities. The study represents the first demonstration of “embryonic” learning in amphibians, and the functions and importance of early olfactory learning for amphibians are discussed.     

An odorant congruent with a colour cue is selectively perceived in an odour mixture

Odour identification can be influenced by colour cues. This study examined the mechanism underlying this colour context effect. We hypothesised that a specific odour component congruent with a colour would be selectively perceived in preference to another odour component in a binary odour mixture. We used a ratio estimation method under two colour conditions, a binary odour mixture (experiment 1) and single chemicals presented individually (experiment 2). Each colour was congruent with one of the odour components. Participants judged the perceived mixture ratio in each odour container on which a colour patch was pasted. An influence of colour was not observed when the odour stimulus did not contain the odour component congruent with the colour (experiment 2); however, the odour component congruent with the colour was perceived as more dominant when the odour stimulus did contain the colour-congruent odorant (experiment 1). This pattern indicates that a colour-congruent odour component is selectively perceived in an odour mixture. This finding suggests that colours can enhance the perceptual representation of the colour-associated component in an odour mixture.     

The discrimination of dog odours by humans

The domestic dog is able to identify both individual conspecifics, i.e. other dogs, and individual interspecifics, e.g. humans, by smell. Whilst humans can recognise individual members of their own species using olfactory information, it is unknown whether they can identify members of any other species in this manner. We examined the ability of humans to identify individual dogs by smell. Twenty-six dog owners were required to smell two blankets, one impregnated with the odour of their own dog, the other impregnated with the odour of an unfamiliar dog. Participants were required to indicate which of the odours smelt the strongest, which smelt the most pleasant, and which of the odours belonged to their own dog. Most of the participants (88.5%) were able to recognise the odour of their own dog. They showed no significant bias, however, in responding which of the odours they thought smelt the strongest or most pleasant. The results indicate that dogs produce odours that are individually distinctive to their owners, and highlight the fact that humans can recognise members of another species using olfactory cues--an ability presumably acquired without conscious effort.     

Developmental differences in memory for cross‐modal associations

Associative learning is critical to normal cognitive development in children. However, young adults typically outperform children on paired‐associate tasks involving visual, verbal and spatial location stimuli. The present experiment investigated cross‐modal odour–place associative memory in children (7–10 years) and young adults (18–24 years). During the study phase, six odours were individually presented and paired with one of 12 spatial locations on a board. During the test phase, participants were presented with the six stimuli individually and were asked to place each stimulus on the correct spatial location. Children committed significantly more errors on the odour–place task than did young adults. However, item recognition memory for the odours or spatial locations involved in the odour–place associative memory task was similar between children and young adults. Therefore, poor odour–place associative memory in children did not result from impaired memory for the individual odours or spatial locations involved in the associations. The results suggest that cross‐modal associative memory is not fully developed in children.     

Acquired hedonic and sensory characteristics of odours: Influence of sweet liker and propylthiouracil taster status

Repeated pairings of novel food-related odours with sweet tastes can result in enduring changes in sweetness of the odour alone, but have less consistent effects on odour liking. Variation in ability to taste propylthiouracil (PROP) might account for this, since PROP supertasters (ST) have been reported both to experience stronger sweetness intensity and to be more likely to dislike sweetness than do PROP nontasters (NT). Alternatively, individual differences in liking for sweetness may transfer to sweet-paired odours independently of PROP sensitivity. To explore this, evaluations of sucrose, saccharin, and PROP solutions were used to classify 92 volunteers as either sweet likers or dislikers and as PROP ST, NT, or medium tasters (MT). Changes in pleasantness of odours that had been paired with the taste of saccharin increased in sweet likers but decreased in dislikers. Odour sweetness increased regardless of PROP taster or sweet liker status. PROP ST rated saccharin as more bitter than did other taster groups and also showed greater increases in acquired bitterness of the saccharin-paired odour. Overall, these data suggest that individual differences in evaluation of saccharin reliably predict subsequent changes in evaluation of saccharin-paired odours, with hedonic changes corresponding with liking for sweet tastes and sensory changes reflecting differences in sensory quality between PROP taster groups.     

Reduced discriminability following perceptual learning with odours

In this experiment, the effects of prior experience on odour perception and discrimination were explored. Participants repeatedly sniffed a mixture composed of two odours, AX, as well as smelling two further odours alone, B and Y. After this training phase, participants were asked to rate the similarity of the odours A and X, B and Y, and a non-exposed pair C and Z. A and X were judged as significantly more alike than the other pairs. Exactly the same pattern emerged on a second test, in which participants were asked to select the odd odour out of sets of three. It was consistently harder for participants to pick the odd odour when the stimuli were drawn from the AX pair (eg A versus AX versus AX). Not only do these findings demonstrate that prior experience can affect odour perception, a finding not predicted by theories of odour perception based solely upon the physiochemical properties of odours, they also suggest that experience can act to selectively decrease discriminability.     

Time of day modulation of conditioned place preference in rats depends on the strain of rat used

In golden hamsters, the expression of a conditioned place preference (CPP) or avoidance (CPA) is regulated in a circadian pattern such that the preference and avoidance are exhibited strongly at the circadian time of prior training, but not at other circadian times. In the rat, reports are conflicting regarding whether time of day learning is evident. We investigated whether this conflict arises because different strains of rat have been used. In this experiment, Long Evans and Wistar rats were trained at a specific circadian time to discriminate between a context paired with food reward and an unpaired context. Animals were then tested for preference at the same or a different circadian time. Long Evans rats showed preference for the paired context at both times tested, whereas Wistar rats showed preference only when training and testing times matched. The results show that time of day learning can be generalized to rats using the Wistar strain.     

Scented memories of literature

This study examined the effects on recall of story details of congruity or incongruity between the hedonic valence of literary texts and odours inhaled while reading them. During the reading session, 24 undergraduates (12 males and 12 females) read two passages involving positive subject matter and two with negative subject matter while sniffing pleasant or unpleasant odours in a within-subject fully counterbalanced design. Subjects rated their experience of each text on eleven 7-point scales. During the test session 48 hours later, subjects read a two-word title associated with each of the passages and inhaled the odour that was paired with it in the reading session. They also rated their experience on six of the scales that had been used during the reading session. Results showed that hedonic congruence between the passage and the odour fostered enhanced recall during the test session. The combination of positive subject matter and positive odour was reflected in more accurate recall of character details, while pairing negative subject matter and negative odour resulted in more accurate recall of setting details. Regression analysis showed that overall recall accuracy was increased by identifying with the characters in the stories and for passages that were found pleasing and personally meaningful. Consistent with the literature on implicit learning involving odours, recall accuracy varied inversely with perceived odour intensity. Implicit learning involving odours and literary passages is therefore fostered by unity in the reading experience.     

Scented memories of literature

This study examined the effects on recall of story details of congruity or incongruity between the hedonic valence of literary texts and odours inhaled while reading them. During the reading session, 24 undergraduates (12 males and 12 females) read two passages involving positive subject matter and two with negative subject matter while sniffing pleasant or unpleasant odours in a within‐subject fully counterbalanced design. Subjects rated their experience of each text on eleven 7‐point scales. During the test session 48 hours later, subjects read a two‐word title associated with each of the passages and inhaled the odour that was paired with it in the reading session. They also rated their experience on six of the scales that had been used during the reading session. Results showed that hedonic congruence between the passage and the odour fostered enhanced recall during the test session. The combination of positive subject matter and positive odour was reflected in more accurate recall of character details, while pairing negative subject matter and negative odour resulted in more accurate recall of setting details. Regression analysis showed that overall recall accuracy was increased by identifying with the characters in the stories and for passages that were found pleasing and personally meaningful. Consistent with the literature on implicit learning involving odours, recall accuracy varied inversely with perceived odour intensity. Implicit learning involving odours and literary passages is therefore fostered by unity in the reading experience.     

Olfactory hedonic context affects both self-report and behavioural indices of palatability

Prior exposure to either a pleasant or unpleasant context may affect later hedonic judgments of a common target stimulus. We explored whether this effect translates into behaviour in the chemical senses. In experiment 1 participants experienced either a pleasant or unpleasant set of odours or pictures, followed by an unfamiliar odour. After self-report hedonic evaluations of the odour, participants were allowed to drink it in solution, followed by a further evaluation of its flavour. Participants reported liking the odour less after smelling pleasant odours and drank less of it too, relative to the unpleasant context. There was no differential context effect for emotive pictures. Experiment 2 replicated these effects, but also included a no-context control. This revealed that the consumption effect was localised to the pleasant olfactory context, whilst contextual effects for liking ratings were primarily localised to the unpleasant olfactory context. In conclusion, hedonic context affects both self-report and behaviour, but not in the same way.     

Are olfactory images sensory in nature?

We investigated the features of olfactory mental images by comparing odour images with perceptual and semantic representations. Participants who were assigned to three groups made similarity judgments about 17 common odours by smelling odours, imagining odours, or on the basis of the meaning of odour source names. In the smelling group, every pair of odours was compared. In the imagining group, imagined odours were compared twice, both before and after associative learning of the odour/name combinations. In the meaning group, the odour source names were compared in terms of general word meanings. Nonmetric multidimensional scaling analysis was applied to each group of similarity data and three-dimensional sensory, mental, and semantic spaces were composed. 17 elements in the mental and semantic spaces were superimposed onto the sensory space by Procrustes rotation. We found that the averaged distances of the 17 elements between the sensory and the mental spaces (either before or after learning) were smaller than those between the sensory and semantic spaces. We suggest that odour images have sensory features, especially after associative learning between perceived odours and their names.     

Choice of extinction-signalling stimuli by rats

Rats in a two-lever situation were exposed to alternating periods of intermittent reinforcement and extinction. Extinction periods were either unsignalled or were signalled by a response-produced stimulus. The signal was sometimes a stimulus paired with food delivery in the reinforcement periods and sometimes a stimulus that occurred only in extinction periods. Both kinds of signal accelerated extinction relative to the unsignalled condition. When the signal was the stimulus paired with food in reinforcement periods, the rats tended to prefer the lever that gave that signal even though the signal accelerated extinction. There was no comparable effect for the stimulus that occurred only in extinction periods; when this signal was contingent on only one of the two levers, the rats either avoided it (Experiment 3) or were indifferent (Experiment 4). It is concluded that a stimulus can be a “secondary reinforcer” as measured by preference, even though it decreases resistance to extinction; the implications are discussed with reference to formal theories of choice behavior.     

The role of pre-exposure to novel food tastes in activity-based conditioned taste avoidance

Rats were given differential exposure to three distinct and novel foods. One of these foods was exposed for 7 days; another for 2 days, and the last was not exposed. Next, half of the rats received six daily sessions in which a compound of the three flavors was followed by opportunities to run in wheels. The other rats received the food compound but without wheel running. On the next day, all rats were given a choice among the three food flavors presented concurrently in separate dishes. When the compound food had been followed by wheel running, rats ate little of the food given no pre-exposure, more of the food given 2 days of pre-exposure, and considerably more of the food given 7 days of pre-exposure. In comparison, rats who did not receive an opportunity to run ate equal and moderate amounts of the three foods. The results suggest that pre-exposure to a food’s taste produces latent inhibition that interferes with conditioned taste avoidance produced by pairing a taste (CS) with wheel running (US).     

Phenomenological differences between familiar and unfamiliar odours

Unfamiliar odours are harder to discriminate than familiar odours. We explored the phenomenal basis of this difference. In experiments la and 1b, participants profiled odour quality for two sets of familiar and unfamiliar odours. In both cases unfamiliar odours were redolent of more odour qualities than familiar stimuli. In experiment 2, participants received (i) a set of familiar and unfamiliar odours and learnt their names, and (ii) a further set of familiar and unfamiliar odours to which they were exposed. Participants then profiled these stimuli as well as a further unexposed set of familiar and unfamiliar odours. Exposure, but not naming, led to a significantly smaller difference between the familiar and unfamiliar stimuli, in terms of their redolence to other odours, when compared to unexposed control stimuli. Unfamiliar exposed odours were also judged as less redolent than unexposed unfamiliar odours. These observations are consistent with a mnemonic basis for odour-quality perception.