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| Odorant Concentration |
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Odorants tend to smell more intense with increasing
vapor phase concentration, but most odorants continue to
elicit the same quality of odor perception across a wide
range of concentrations. The exceptions to this rule are
notable, however. About 8% of the odorants surveyed in
an archive of flavor and fragrance chemicals were
reported to change in their odor quality with
concentration (Arctander, 1994). The differences in odor
can be striking: for example, beta-ionone is reported to
smell like violets at low concentrations and like cedar
at higher concentrations (Dravnieks, 1985).
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We studied the patterns of 2-DG uptake evoked by five
aliphatic compounds differing in functional groups
(Johnson and Leon, 2000a). Three of the compounds
(valeric acid, 1-pentanol, and methyl valerate) were not
reported by humans to change in perceived odor quality
with stimulus concentration, whereas the other two
compounds (2-hexanone and pentanal) were reported to
evoke distinct odor perceptions at high and low
concentrations. When we expressed our data in units of z
scores to illustrate the pattern of uptake independently
of the amount of uptake, we found that the evoked
activity patterns were constant with concentration for
the three odorants that displayed perceptual constancy,
and that the activity patterns differed with
concentration for the two odorants that differed in
perceived odor at high and low concentrations (Johnson
and Leon, 2000a). This finding represents yet another
correlation between activity patterns as measured by
2-DG uptake and perceived odor, assuming that rats
perceive the odors of these compounds in a manner
similar to humans. These data also suggest that odor
quality perception may be coded in terms of the relative
pattern of activity in particular neurons compared to
the overall level of activity across the entire
olfactory bulb. It may be the case that the perception
of odor changes when concentration is increased for some
odorants because increasing the concentration of the
target odorant also increases odorant contaminants that
affect both neural pattern and odor perception. In the
case of pentanal, the acid contaminant in pentanal,
pentanoic acid, could reasonably account for the
observed changes in neural response and odor perception
(Johnson et al., 2004).
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The z-score activity pattern
evoked by 2-hexanone changed with odorant concentration,
whereas the z-score pattern evoked by valeric acid was
constant across different concentrations.
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When we expressed our data in units allowing us to see
the amount of 2-DG uptake in addition to the pattern of
uptake, increasing stimulus concentration was associated
with increased 2-DG uptake in most cases (Johnson and
Leon, 2000a). The general increase in activity gave the
impression that individual activity foci grew in size,
recruiting additional glomeruli at any given threshold
value of activity. This pattern is similar to what has
been found using other methods of monitoring glomerular
activity (Guthrie and Gall, 1995; Rubin and Katz, 1999;
Meister and Bonhoeffer, 2001; Wachowiak and Cohen,
2001). The major exception to this rule of increasing
activity with increasing odorant concentration involved
high concentrations of valeric acid, where 2-DG uptake
decreased in all response modules at the highest
concentration tested (Johnson and Leon, 2000a). Rats
withhold their inspiration at high concentrations of
valeric acid, which is an irritant detected by the
trigeminal nerve (Alarie, 1973). Decreased sniffing of
valeric acid at high vapor concentrations could explain
the phenomenon of decreased 2-DG uptake at these high
stimulus concentrations.
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When expressed in terms of
glomerular layer 2-DG uptake divided by subependymal
zone 2-DG uptake, activity was seen to increase with
concentration up to 12 parts per million of valeric
acid, followed by a decline that may reflect decreased
sniffing of the irritating odorant.
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