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Odors elicit a variety of behavioral responses from Drosophila via a relatively simple, but sensitive, olfactory system. An increasing number of mutants have been found to be defective in olfactory function. Genetic and molecular analyses of the Drosophila olfactory system have identified some of its molecular components, and have revealed some principles of its function and organization.
1. Several members of the family of heterotrimeric signal transducing GTP-binding proteins have been identified in the olfactory epithelium of vertebrates by biochemical and molecular cloning techniques. 2. Biochemical and electrophysiological evidence indicates that one or more GTP-binding proteins mediate many olfactory responses by coupling stimulus receptors to second messenger systems. 3. Although GTP-binding proteins may function in additional transduction events, a novel GTP-binding protein, expressed only in olfactory neurons, may mediate stimulus activation of adenylate cyclase in olfactory cilia.
Female insects release sex-pheromones which attract their conspecific males. These pheromones are detected through a distinct male-specific olfactory subsystem which resides at the first stage of olfactory processing, and consists of receptor, local and projection (relay) neurons. When male insects were stimulated by female sexpheromones, some projection neurons could distinguish between different pheromones, following input and code stimulus intensity. Presented here, is a simple biophysical model that described characteristic bursting responses observed for projection neurons. The bursting behavior of the model resulted from a particular cellular mechanism and specific network architecture. At the neuron level, a rapidly activating and slowly inactivating low-threshold calcium channel provided depolarizing current for bursting, while at the network level, inhibitory neurons implementing dis-inhibition which triggered this calcium channel. Also, the network architecture provided a mechanism by which certain projection neurons coded temporal input and stimulus intensity.
Olfactory stimuli (odorants) are detected and recognized by binding to receptors belonging to the G-protein-coupled receptor superfamily. The binding of odorants to some receptors stimulates the activity of an odorant-sensitive phospholipase C (PLC) thereby generating the second messengers inositol 1,4,5-trisphosphate (IP3) and diacylglycerol (DAG). IP3 plays a key role in membrane depolarization by binding to a receptor that is itself a cation channel. The formation of DAG is expected to stimulate the activity of protein kinase C (PKC). PKC, together with G-protein-coupled receptor kinases, mediates signal termination by phosphorylation of odorant receptors and possibly other substrates. This review summarizes recent evidence regarding the role of phosphoinositide-derived second messengers in the molecular events underlying olfactory signaling. In addition, the role of calcium as a "third messenger" that provides a mechanism for interaction between phosphoinositide second messengers and components of the cyclic AMP signaling pathway is also discussed.
Retinal projections to several telencephalic structures have been demonstrated in a wide range of mammalian species following intraocular injections of tritiated amino acids and cholera toxin subunit-B conjugated to horseradish peroxidase. Since these regions are also innervated by olfactory fibers, we investigated the distribution of convergent projections using simultaneous injections of different anterograde tracers in the eye and olfactory bulbs. Convergent projections from the retina and from the olfactory bulbs were observed in the piriform cortex, olfactory tubercle, the cortical region of the medial amygdala, lateral hypothalamus, and the bed nucleus of the stria terminalis. A few retinal fibers also invade the nucleus of the lateral olfactory tract, the bed nucleus of the accessory olfactory bulb and the diagonal band of Broca. Injections of retrograde tracers in the medial amygdala, the bed nucleus or the lateral hypothalamus shows that the visuo-olfactory convergence mainly involves projections originating from the accessory olfactory bulb, and to a lesser extent from the ventromedial region of the main olfactory bulb. Fewer than 20 retinotelencephalic ganglion cells were identified in the retina, mainly located contralateral to the injection site. Ganglion cells were medium sized and possessed two long slender opposing dendrites. These retinal and olfactory projections could provide an anatomical substrate for the modulation of gonadotropin hormone levels and the olfactory influence on light mediated rhythms related to reproductive physiology.
We have isolated nine putative odorant receptor genes from the chick, named COR1 to COR9, that belong to the large multigene family of olfactory G protein-coupled receptors found in the fish, rat, mouse, dog, and human. By combining genomic DNA blot analysis, low stringency library screenings, and several PCR analyses, we were able to detect approximately 20 COR genes in the chick genome highly related to COR1-9. By in situ hybridization of newborn and adult, COR expression was detected only in the olfactory epithelium, and exhibited a random spatial distribution. During development, COR expression was observed as early as embryonic stage E5. Different levels of gene expression were observed for the COR1-9 genes: at E5, COR1-6 expression was high compared to the expression of COR7, COR8, and COR9. Surprisingly, at E5, a row of COR1-6 positive cells probably associated with the olfactory nerve extended outside the olfactory placode, reaching the anterior pole of the developing forebrain. These results suggest that, in addition to their role as putative odorant receptors, some COR may play a role in the development of the avian olfactory system.
Studies on the properties of olfactory receptors and of the olfactory glomeruli indicate that there is spatial segregation of response to particular characteristics of odorant molecules at the input level of the olfactory bulb. Existing anatomical information and studies of synaptic mechanisms in the olfactory bulb suggest that the bulb circuitry might act as a contrast detection mechanism analyzing a spatially organized input. Recent electrophysiological studies have supported this idea. Extracellular recordings have shown that the similarity between responses of cell pairs to the same stimulus odor depend upon the distance between those cells. Intracellular recordings from mitral and tufted cells have shown spatially organized excitatory and inhibitory responses to localized electrical stimulation of the input layer of the bulb. Some of the major interneurons of the olfactory bulb have also been identified during odor and localized electrical stimulation. These recordings are also consistent with a spatially based organization.
The relationship between odor identification and cognition has not been previously well characterized. The neuroanatomy of the olfactory system and the frequent finding of olfactory dysfunction in neurodegenerative diseases suggest a likely relationship between odor identification and memory, language, and executive functioning, though previous studies have often failed to demonstrate the expected relationship. The current study examined this relationship in across a continuum of ability levels (N=100). Strongest correlations were found between odor identification and language, most aspects of memory, and a measure of general cognitive functioning. Significant but more modest correlations were seen between odor identification and attention, motor, visuospatial, and executive functions. A regression analysis revealed language as the only significant predictor of olfactory performance. These findings suggest that odor identification is most closely associated with other measures of temporo-limbic functioning. The implications of these findings, particularly in consideration of the assessment of older adults, are discussed.
Odorant receptor genes are expressed not only in the nose but also in testes, where they have been hypothesized to play a role in sperm chemotaxis. New data demonstrate that human odorant receptor hOR 17-4 may play similar roles in both tissues, lending support to the idea that chemical attraction is important for reproduction.
Humans use three classes of photoreceptor to span the visible spectrum, but smell relies on hundreds of distinct classes of olfactory receptor neuron. Even the simple fruitfly has around 50 classes of olfactory receptor neuron. Two new studies map the projections of the great majority of these neurons into stereotyped positions in the fly brain, giving us an almost complete atlas of olfactory information transfer.
Lateralization of function is a well-known phenomenon in humans. The two hemispheres of the human brain are functionally specialized such that certain cognitive skills, such as language or musical ability, conspecific recognition, and even emotional responses, are mediated by one hemisphere more than the other [1, 2]. Studies over the past 30 years suggest that lateralization occurs in other vertebrate species as well [3-11]. In general, lateralization is observed in different sensory modalities in humans as well as vertebrates, and there are interesting parallels (reviewed in [12]). However, little is known about functional asymmetry in invertebrates [13, 14] and there is only one investigation in insects [15]. Here we show, for the first time, that the honeybee Apis mellifera displays a clear laterality in responding to learned odors. By training honeybees on two different versions of the well-known proboscis extension reflex (PER) paradigm [16, 17], we demonstrate that bees respond to odors better when they are trained through their right antenna. To our knowledge, this is the first demonstration of asymmetrical learning performance in an insect.
In identifying the volatiles that insects use to locate suitable host plants, electrophysiological recordings of olfactory responses to plant volatiles may give important information. However, divergent results may be obtained with different recording techniques. To illustrate these differences, the results of a previous investigation using single cell recordings linked to a gas chromatograph (SCR-GC) are compared with the results obtained with an electroantennogram linked to a gas chromatograph (EAG-GC). Testing insects of the same species (Hylobius abietis) for the same test sample, 30 potent volatile compounds were identified by SCR-GC and 18 by EAG-GC. Of the 34 different compounds, 14 were identified by both techniques. Furthermore, when the same compound elicited detectable responses by both techniques, the response strength was usually not the same relative to the strongest response recorded by each technique. This shows that both EAG-GC and SCR-GC are important techniques in the identification of potent plant volatiles for insects. However, by using SCR-GC more information was obtained, information that can be crucial for understanding the insect-plant relationship.
The olfactory system has generated considerable interest in recent years, mainly focused on receptor genes and early olfactory processing. In this issue of Neuron, Mori et al. focus centrally, providing evidence for slow- and fast-wave states in olfactory cortex that appear to gate the inflow of information underlying conscious smell perception.
OBJECTIVE: To see if nasal peak inspiratory flow rate and subjective sense of smell had any correlation with olfactory thresholds. STUDY DESIGN AND SETTING: A cohort study of 186 normal volunteers was recruited from among staff and visitors at a university hospital. Olfactory thresholds were detected for each subject (103 with eucalyptol and 83 with phenethyl alcohol), along with nasal peak inspiratory flow (PIFR). Subjective sense of smell, along with nasal symptoms, mood, and alertness, were recorded on visual analogue scores. RESULTS: Subjective perception of smell had no correlation with olfactory thresholds detected (P = 0.4057) and the other subjective measures also had no correlation. There was a significant relationship of PIFR to thresholds in the group tested with phenethyl alcohol (P = 0.002). CONCLUSION: As with the sensation of nasal patency, the self-assessment of a subject's sense of smell has poor correlation with their actual olfactory ability. SIGNIFICANCE: A patient's history cannot be relied upon when determining their olfactory ability and formal testing should be performed. EBM RATING: A-1b.
Means to boost food intake of the elderly (n = 60, age range 61-86 years) varying in olfactory and mood status were studied during home-use of cold cuts. Enhancement group (n = 28) received regular smoked ham and smoked ham flavored with extra strong smoke aroma or with pepper mix, and Variety group (n = 32) received four different cold cuts: smoked ham, cooked ham, pepper ham and bologna. Both groups participated in two successive home-use periods, each over 6 days. Before and after home-use, the cold cuts were rated for odor and flavor intensity and liking, and in home-use only for liking. Cold cuts, 600 g per period, were to be consumed ad libitum, and the returned left-overs were weighed. Scandinavian Odor Identification Test and Profile of Mood States (bi-polar) were used in dividing subjects into normosmics and hyposmics (SOIT, cut-off score: 11) and into mood groups (POMS, depressed-elated, cut-off score: median). Enhancement group liked both the regular and the ham enhanced with pepper mix similarly, while added smoke aroma decreased liking. Regardless of the olfactory capabilities, the subjects >74 years of age were less responsive to the flavor changes in their ratings than those aged 61-74 years. Poor olfactory status increased the intakes regardless of the flavor level or type of cold cuts. Path analysis suggested that in the Enhancement group, poor health, depressed mood and high age predicted a poor olfactory performance which, along with ratings of liking, increased the consumed amounts of cold cuts.
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Large numbers of the snail Terebralia palustris (Linnaeus) (Potamididae; Gastropoda) are frequently observed feeding in a cluster on a single fallen mangrove leaf, yet none are present on leaves nearby. Consequently, we investigated the food-finding ability of T. palustris in a Kenyan mangrove forest using field experiments. We estimated the attractive effect of different cues and analysed the paths (video-recorded) of snails when approaching a food-related odour. This intertidal snail can potentially use both air-borne and water-borne odours to locate food. T. palustris is attracted to conspecifics feeding on leaves, while intact leaves as well as non-feeding snails are not attractive. Moreover, the guiding stimulus appears to be compounds released when the leaves are damaged.T. palustris also seems able to discriminate between different food items; it is more strongly attracted to green mangrove leaves than senescent or fallen ones or mangrove propagules, probably because green leaves release a greater amount of attractive cues.Feeding snails thus recruit more snails to feed on the same leaf. The ecological implications of this behaviour are discussed: a large number of snails on the same leaf counteracts the ability of crabs to remove the leaf being eaten by the snails.