Sensory Systems/Olfactory System
- 1 Introduction
- 2 Sensory Organs
- 3 Sensory Organ Components
- 4 Signal Processing
- 5 References
Probably the oldest sensory system in the nature, the olfactory system concerns the sense of smell. The olfactory system is physiologically strongly related to the gustatory system, so that the two are often examined together. Complex flavors require both taste and smell sensation to be recognized. Consequently, food may taste “different” if the sense of smell does not work properly (e.g. head cold).
Generally the two systems are classified as visceral sense because of their close association with gastrointestinal function. They are also of central importance while speaking of emotional and sexual functions.
Both taste and smell receptors are chemoreceptors that are stimulated by molecules soluted respectively in mucus or saliva. However these two senses are anatomically quite different. While smell receptors are distance receptors that do not have any connection to the thalamus, receptors pass up the brainstem to the thalamus and project to the postcentral gyrus along with those for touch and pressure sensibility for the mouth.
In this article we will first focus on the organs composing the olfactory system, then we will characterize them in order to understand their functionality and we will end explaining the transduction of the signal and the commercial application such as the eNose.
In vertebrates the main olfactory system detects odorants that are inhaled through the nose where they come to contact with the olfactory epithelium, which contains the olfactory receptors.
Olfactory sensitivity is directly proportional to the area in the nasal cavity near the septum reserved to the olfactory mucous membrane, which is the region where the olfactory receptor cells are located. The extent of this area is a specific between animals species. In dogs, for example, the sense of smell is highly developed and the area covered by this membrane is about 75 – 150 cm2; these animals are called macrosmatic animals. Differently in humans the olfactory mucous membrane cover an area about 3 – 5 cm2, thus they are known as microsmatic animals.
In humans there are about 10 million olfactory cells, each of which have 350 different receptor types composing the olfactory mucous membrane. The 350 different receptors are characteristic for only one odorant type. The bond with one odorant molecule starts a molecular chain reaction, which transforms the chemical perception into an electrical signal.
The electrical signal proceeds through the olfactory nerve’s axons to the olfactory bulbs. In this region there are between 1000 and 2000 glomerular cells which combine and interpret the potentials coming from different receptors. This way it is possible to unequivocally characterise e.g. the coffee aroma, which is composed by about 650 different odorants. Humans can distinguish between about 10.000 odors.
The signal then goes forth to the olfactory cortex where it will be recognized and compared with known odorants (i.e. olfactory memory) involving also an emotional response to the olfactory stimuli.
It is also interesting to note that the human genome has about 600 – 700 genes (~2% of the complete genome) specialized in characterizing the olfactory receptors, but only 350 are still used to build the olfactory system. This is a proof of the evolution change in the necessity of humans in using the olfaction.
Sensory Organ Components
Similar to other sensory modalities, olfactory information must be transmitted from peripheral olfactory structures, like the olfactory epithelium, to more central structures, meaning the olfactory bulb and cortex. The specific stimuli has to be integrated, detected and transmitted to the brain in order to reach sensory consciousness. However the olfactory system is different from other sensory systems in three fundamental ways as depicted in the book of Paxianos G. and Mai J.K., "The human Nervous System".
- Olfactory receptor neurons are continuously replaced by mitotic division of the basal cells of the olfactory epithelium. The motivation of this is the high vulnerability of the neurons, which are directly exposed to the environment.
- Because of phylogenetic relationship, olfactory sensory activity is transferred directly fro the olfactory bulb to the olfactory cortex, without a thalamic relay.
- Neural integration and analysis of olfactory stimuli may not involve topographic organization beyond the olfactory bulb, meaning that spatial or frequency axis are not needed to project the signal.
Olfactory Mucous Membrane
The olfactory mucous membrane contain the olfactory receptor cells and in humans it covers an area about 3 – 5 cm^2 in the roof of the nasal cavity near the septum. Because the receptors are continuously regenerated it contains both the supporting cells and progenitors cells of the olfactory receptors. Interspersed between these cells are 10 – 20 millions receptor cells.
Olfactory receptors are infect neurons with a short and thick dendrites. Their extended end is called an olfactory rod, from which cilia project to the surface of the mucus. These neurons have a length of 2 micrometers and have between 10 and 20 cilia of diameter about 0.1 micrometers.
The axons of the olfactory receptor neurons go through the cribriform plate of the ethmoid bone and enter the olfactory bulb. This passage is in absolute the most sensitive of the olfactory system; the damage of the cribriform plate (e.g. breaking the nasal septum) can imply the destruction of the axons compromising the sense of smell.
A further particularity of the mucous membrane is that with a period of a few weeks it is completely renewed.
In humans the olfactory bulb is located anteriorly with respect to the cerebral hemisphere and remain connected to it only by a long olfactory stalk. Furthermore in mammals it is separated into layers and consist of a concentric lamina structure with well-defined neuronal somata and synaptic neuropil.
After passing the cribriform plate the olfactory nerve fibers ramify in the most superficial layer (olfactory nerve layer). When these axons reach the olfactory bulb the layer gets thicker and they terminate in the primary dendrites of the mitral cells and tufted cells forming in this way the complex globular synapses called olfactory glomeruli. Both these cells send other axons to the olfactory cortex and appear to have the same functionality but in fact tufted cells are smaller and consequently have also smaller axons.
The axons from several thousand receptor neurons coverage on one or two glomeruli in a corresponding zone of the olfactory bulb; this suggest that the glomeruli are the unit structures for the olfactory discrimination.
In order to avoid threshold problems in addition to mitral and tufted cells, the olfactory bulb contains also two type of cells with inhibitory properties: periglomerular cells and granule cells. The first will connect two different glomeruli, the second, without using any axons, build a reciprocal synapses with the lateral dendrites of the mitral and tufted cells. By releasing GABA the granule cell on the one side of these synapse are able to inhibits the mitral (or tufted) cells, while on the other side of the synapses the mitral (or tufted) cells are able to excite the granule cells by releasing glutamate. Nowadays about 8.000 glomeruli and 40.000 mitral cells have been counted in young adults. Unfortunately this huge number of cells decrease progressively with the age compromising the structural integrity of the different layers.
The axons of the mitral and tufted cells pass through the granule layer, the intermediate olfactory stria and the lateral olfactory stria to the olfactory cortex. This tract forms in humans the bulk of the olfactory peduncle. As depicted in the book of Paxianos G. and Mai J.K., "The human Nervous System", the primary olfactory cortical areas can be easily described by a simple structure composed of three layers: a broad plexiform layer (first layer); a compact pyramidal cell somata layer (second layer) and a deeper layer composed by both pyramidal and nonpyramidal cells (third layer). Furthermore, in contrast to the olfactory bulb, only a little spatial encoding can be observed; “that is, small areas of the olfactory bulb virtually project the entire olfactory cortex, and small areas of the cortex receive fibers from virtually the entire olfactory bulb” .
In general the olfactory tract can be divided in five major regions of the cerebrum: Anterior olfactory nucleus, the olfactory tubercle, the piriform cortex, Anterior cortical nucleus of the amygdala and the entorhinal cortex.Olfactory information is transmitted from primary olfactory cortex to several other parts of the forebrain, including orbital cortex, amigdala, hippocampus, central striatum, hypothalamus and mediodorsal thalamus.
Interesting is also to note that in humans, the piriform cortex can be activated by sniffing, whereas the to activate the lateral and the anterior orbitofrontal gyri of the frontal lobe only the smell is required. This is possible because in general the orbitofrontal activation is grater on the right side than the left side, this directly imply an asymmetry in the corticals reception of the olfaction. A further implication of the emotional response to olfactory stimuli as olfactory memories can be assigned to the fibers projection to the amigdala of the entorhinal cortex.
A good and complete description of the substructure of the olfactory cortex can be found in the book of Paxianos G. and Mai J.K., "The human Nervous System".
|Substance||mg/L of Ari|
|Oil of peppermint||0.02|
Only substances which comes in contact with the olfactory epithelium can be excite the olfactory receptors. The right table shows some threshold for some representative substances. These values give an impression of the huge sensitivity of the olfactory receptors.
It is remarkable that humans can recognize more than 10'000 different odors but they should at least differ about the 30% before they can be distinguished. Compared to the visual system, such precision would mean a 1% change in light intensity, where as compared to hearing the direction perception may be indicated by the slight difference in the time of arrival of odoriferous molecules in the two nostrils . It is amazing how the same number of carbon atoms (normally between 3 and 20) in odors molecules can leads to different odors just by slightly change in the structural configuration.
An interesting feature of the olfactory system is how a simple sense organ that apparently lacks a high degree of complexity can mediate discrimination of more than 10'000 different odors. On the one hand this is made possible by the huge number of different odorant receptor. The gene family for the olfactory receptor is infect the largest family studied so far in mammals. On the other hand the neural net of the olfactory system’s provide with their 1800 glomeruli a large two dimensional map in the olfactory bulb that is unique to each odorant. In addition, the extracellular field potential in each glomerulus oscillates, and the granule cells appear to regulate the frequency of the oscillation. The exact function of the oscillation is unknown, but it probably also helps to focus the olfactory signal reaching the cortex .
Olfaction, as described in the research of R. Haddad et al., consists of a set of transforms from physical space of odorant molecules (olfactory physicochemical space), through a neural space of information processing (olfactory neural space), into a perceptual space of smell (olfactory perceptual space). The rules of these transforms depend on obtaining valid metrics for each of those spaces.
Olfactory perceptual space
As the perceptual space represent the “input” of the smell measurement, it’s aim is to describe the odors in the most simple possible way. Odor are infect ordered so that their reciprocal distance in space confers them similarity. This mean that odors the more two odors are near each other in this space the more are they expected to be similar. This space is thus defined by so called perceptual axes characterized by some arbitrarily chosen “unit” odors.
Olfactory neural space
As suggested by its name the neural spaces are generated from neural responses. This gives rise to an extensive database of odorant-induced activity, which can be used to formulate an olfactory space where the concept of similarity serves as a guiding principal. Using this procedure different odorant are than expected to be similar if they generate a similar neuronal response. This database can be navigated at the Glomerular Activity Response Archive .
Olfactory physicochemical space
The need of identify the molecular encryption of the biological interaction, make the physicochemical space the most complex one of the olfactory space described so far. R. Haddad suggest that one possibility is to span this space would to represent each odorant by a very large number of molecular descriptors by use either a variance metric or a distance metric. In his first description single odorants may have many physicochemical features and one expect these feature to present themselves at various probabilities within the world of molecules that have a smell. In such metric the orthogonal basis generated from the description of the odorant leads to represent each odorant by a single value. While in the second, the metric represents each odorant with a vector of 1664 values, on the basis of Euclidean distances between odorants in the 1664 physicochemical space. Whereas the first metric enabled the prediction of perceptual attributes, the second enabled the prediction of odorant-induced neuronal response patterns.
Electronic measurement of odors
Nowadays odors can be measured electronically in a huge amount of different way, some examples are: mass spectrography, gas chromatography, raman spectra and most recently electronic nose. In general they assume that different olfactory receptors have different affinities to specific molecular physicochemical properties, and that the different activation of these receptors gives rise to a spatio-temporal pattern of activity that reflects odors.
eNoses are analytic devices for mimicking the principle of biological olfaction that have as main component an array of non specific chemical sensors. Combining electronics, path recognition and modern technology, the eNoses uses gas sensors to translate the chemical signal into an electrical signal when an volatile odorant from a sample reaches the gas sensor array. Usually the pattern recognition is used to perform either the quantitative or the qualitative identification. In order to reproduce the olfactory epithelium a gas sensor array is sealed in a chamber of the eNose. A cross-sensitive chemical sensors will than act as olfactory neuron transferring the odor information from a chemical into an electric form similar to the one process which occur in the olfactory bulb where the signal is integrated and enhanced. The information is than elaborated by an artificial neuronal network, which provide coding, processing and storage. The gas sensor array transforms odor information from the sample space into a measurement space. This is a key procedure for information processing within an eNose. Gas sensors with different transduction principles and different fabrication techniques provide various ways to obtain odor information. Commercially a lot of different sensor types are available the most frequently used sensor types include metal oxide semiconductors (MOS), quartz crystal microbalances (QCM), conducting polymers (CP) and surface acoustic wave (SAW) sensors. A big influence in the choice of the sensor is made by the fast response, reversibility, repeatability and high sensitivity of the sensor. While constructing the sensor array for a eNose the sensors are selected to be cross-selective to different odors, such that their sensitivity is overlapped with the same odor, to make the most of type-limited sensors for obtaining adequate odor information. In general the amount of raw data generated from the array of sensor’s is huge, so that the information has to be transferred from a high dimensional space into a lower one. Pattern recognition are then needed to encode the signal into a so called classification space. Both are important and necessary for designing a powerful information processing algorithm and constructing an array with high quality gas sensors. Many pattern recognition methods have been introduced into eNose, including parameterized and non-parameterized multivariate statistical methods. Artificial neural network have various significant advantages: (i) Self-adaptive, (ii) capability of error tolerance and generalization suitable for treating the problems (iii) parallel processing and distributed storage.
- Schmidt, Lang (2007). "Ohysiologie des Menschen", Soringer, 30. Auflage.
- Faller A., Schünke M. (2008). "Der Körper des Menschen", Thieme, 15. Auflage.
- Paxianos G., Mai J.K. (2004). "The human Nervous System", Elsevier accademic press, 2nd Edition.
- William. "Review of Medial Physiology", Lange, 22th Edition.
- Haddad R. ed al (2008). "Measuring smells", Elsevier Ltd, 18:438-444
- Mamlouk A.M., Martinez T. (2004). "One dimensions of the olfatory perception space", Elsevier B.V.
- >Guang L ed al (2009), "Progress in bionic information processing techniques for an electronic nose based on olfactory models", Chinese Science Bulletin, 54(4)521-53Z