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Figure 2—figure supplement 1. Reliable delivery of odor concentration series and concentration-dependent effects on breathing. ; (A) Odor delivery was rapid and reliable across a 30-fold range of concentration. Traces are photoionization detector (PID) output overlaid for 12 trials each of 0.03, 0.1, 0.3 and 1% dilutions of ethyl butyrate in mineral oil. (B) Odor concentration series diluted in mineral oil reliably produced corresponding PID output separated by 0.5 log10 steps. PID output was monitored at final valve exhaust in 8/9 experiments (n = 8). The dashed line shows nominal dilution. (C) Breath period measured immediately after odor delivery (filled circles) and coefficient of variation over four breaths after odor delivery (unfilled circles) as a function of stimulus concentration (n = 12, i.e. 6 experiments x 2 different odorants). Breath period increases systematically with concentration but absolute difference in period is

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Figure 2—figure supplement 2. The mOT neurons were activated by different odor stimulations. ; (A) Schematic of the fiber photometry recording from the mOT neurons. (B) Representative traces of GCaMP fluorescence in response to odor and mineral oil stimulations for the GCaMP6m-expressing C57BL/6 mice.

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Figure 1—figure supplement 2. M71 transgenic mice fail to detect acetophenone in a go/no go operant conditioning task. ; (A) Wild-type mice consistently discriminate acetophenone from its diluent mineral oil (black lines). Wild-type mice also discriminate between other pairs of odorants (ethyl acetate versus mineral oil, citronellol, or carvone, dotted lines). (B) M71 transgenic mice consistently fail to discriminate acetophenone from mineral oil, but readily discriminate between the other pairs of odorants.

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Figure 1. The ability of M71 transgenic mice to detect acetophenone is task-dependent. ; (A, B) In a go/no go operant conditioning task, M71 transgenic mice fail to discriminate acetophenone from mineral oil (left panels). In contrast, M71 transgenic mice readily discriminate other pairs of odorants (ethyl acetate vs. mineral oil, citronellol, or carvone, right panels). (A) Original results reported in Fleischmann et al. (2008). (B) Repeat experiment with an additional cohort of mice. Thin lines: learning curves for individual mice. Thick lines: averaged learning curves. Error bars: 95% CI of the mean. (C) Sniff adaptation: schematic of the experimental configuration. (D) Example sniff traces during first 3 (1st, 2nd, and 3rd) presentations of hexanone (shaded area) from a control mouse. Lighter colored traces signify later presentations. ‘FV’ trace shows opening of final valve directing odorized air to the mouse, ‘flow’ trace shows the output from the olfactometer, and ‘PID’ trace shows signal evoked by odorized air from a photo-ionization detector. (E) Example moving averages of instantaneous sniff frequency during first 3 presentations of hexanone (window = 500 ms, plotted against leading edge). Black traces: controls, red traces: M71 transgenic mice. (F, G) Mean instantaneous sniff frequency responses to first vs. the average of the 2nd and 3rd presentation of an odor for control (black, F) and M71 transgenic (red, G) mice. Pooled non-acetophenone odorants: hexanone, ethyl acetate, heptanal, and an odor mixture. Lighter colors: individual trials, thick lines: averages. Error bars: SD. Black dotted lines on the M71 plots show the means for the corresponding data from controls.

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Figure 1. Odor responses in piriform cortex. ; (A) Schematic of experimental setup. (B) Left, Coronal brain section showing region of anterior piriform cortex where recordings were obtained. Multielectrode probes were painted with DiI to mark recording locations (red, DiI; green, NeuroTrace). DiI in the left hemisphere is from a separate recording. Right, higher magnification image of boxed area on left, superimposed with a schematic at scale of a 32-channel probe. Numerals indicate piriform layers. Note the electrode tip spans layer II. (C) Dorso-ventral (DV) distribution of 459 isolated cells (n = 9 experiments) determined by measuring the center of mass of each unit’s waveform amplitudes recorded on different channels across the probe. (D) Population activity and timing of odor delivery. One-second odor pulses (black bar, isoamyl acetate) were triggered on exhalation. Blue shaded region indicates first full respiration cycle after odor onset. Population raster plots below display spiking activity of 33 simultaneously recorded cells sorted by estimated DV position within layer II. (E) Inhalation-aligned raster plots showing responses of three simultaneously recorded cells to three different odor stimuli (0.3% v./v.). Gray shading indicates odor delivery. Blue shading indicates respiration cycle. Cells displayed varying degrees of odor specificity, activation or suppression, and temporal precision. (F–G) Population responses in a representative experiment with 48 simultaneously recorded neurons sorted by relative DV location responding to six different monomolecular odorants (0.3%). (F). Firing rates during the first respiration cycle after odor onset. A mineral oil control is shown on left. (G). Same data expressed using a response index where cells with response index values of −1 (blue) and 1 (red) are unambiguously suppressed or activated, respectively. Odors: et. bu., ethyl butyrate; 2-hxn., 2-hexanone; is. ac., isoamyl acetate; hex., hexanal; et. ti., ethyl tiglate; et. ac., ethyl acetate. (H–J) Percent cells responding to different numbers of odorants with significantly increased spiking (H), decreased spiking (I), or mixed polarity responses (J). Filled circles are mean ± s.e.m. (n = 9). The dashed line denotes the expected distribution when cell identities are shuffled; shaded area, 5th–95th percentiles.

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Figure 1—figure supplement 3. Properties of Activated and Suppressed cells. ; (A) PSTHs for six example cells demonstrating exclusively odor-evoked activation (cells 2, 3, and 5, red traces) or odor-evoked suppression (cells 1, 4, and 6, blue traces). Response to mineral oil for each cell is overlaid in gray. (B) Example thresholded response map for a representative experiment with 73 cells showing consistent response polarity across odors (left). Example cells are indicated on the left. Many more mixed polarity responses are expected if responses are randomly assigned to cells within an odor (right). (C) ‘Activated’ (n = 87/459) and ‘Suppressed’ (n = 132/459) cells were identified as having exclusively significant activation or suppression to at least one of six odors. Spontaneous firing was measured during intertrial intervals. Activated cells had lower spontaneous firing rates than Suppressed cells (Activated: mean ± st. dev; 3.07 ± 3.90 Hz, n = 87; Suppressed: 5.16 ± 5.35 Hz; n = 132 p

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Figure 4—figure supplement 1. Odorant concentration scales with nominal dilution and breath period is independent of odorant concentration. ; (a) Photoionization detector (PID) traces for Hexanone at 1:10,000, 1:1,000, and 1:100 vol./vol. dilution in mineral oil. Four individual trails and average trace across the four trials are shown. Shaded area: SD of the mean. (b) Coefficient of variation of the mean PID signal after odor onset, averaged across 14 trials. Error bar: SD of the mean. (c) Mean PID signal normalized to 1:10,000 vol./vol. dilution, averaged across 14 trials. Red dotted line shows the expected output from nominal dilutions. Error bar: SD of the mean. (d) Breath period of mice before and after exposure to odorants at different concentrations. Error bar: SD of the mean.

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Figure 9. Dynamics of cortical inhibition indicates their role in shaping piriform responses. ; (A) Schematic of experiment. Optogenetic identification of ChR2-expressing neurons in piriform cortex of VGAT-ChR2-EYFP transgenic mice. (B) Coronal section of anterior piriform cortex of VGAT-ChR2-EYFP mouse showing dense innervation of layer II/III, consistent with feedback inhibition (yellow, anti-GFP; blue, neurotrace). Scale bar, 100 μm. (C) One second laser pulses (2.5 mW/mm2) evoke sustained responses in presumptive ChR2-expressing neurons (blue raster plots) but sustained suppression in most other cells (black). Scale bar, 50 μV, 1 ms. (D) PSTHs and raster plots for three VGAT+ cells responding to mineral oil control or three different odors. (E) Response matrix for all recorded VGAT+ cells (n = 35) responding to six different odors. (F) Tuning for all recorded VGAT+ cells. Responses tended to be positive and broadly-tuned. (G–H) Average PSTHs for all VGAT+ cell responses (lines and shading are mean ± s.e.m; n = 70, 35 cells x 2 odors) at four concentrations. (G) PSTHs shown on a short time scale. Stronger and earlier peaks occur at higher concentrations. (H) Average VGAT+ PSTHs overlaid with the bootstrap confidence intervals (gray bars) for the first and second peaks of the piriform cortex response latency distributions from Figure 3.

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Figure 8. ; H2be is regulated by activity. (A,B) Effects of unilateral naris occlusion (UNO) on H2BE level in the MOE. (A) Representative images of FLAG-H2BE and Olfr733 colocalization (boxed regions magnified, right; Olfr733+ neurons, arrowheads). (B) Distributions (main) and averages (inset) of H2BE level within nuclei of randomly-sampled (main; inset, left; n = 200) or Olfr733+ (inset, right; n = 10) neurons on the two sides of the MOE. (C) Effects of UNO on intrinsic GAP43-mCherry fluorescence in glomeruli (arrowheads) within the OB of an H2be-KO heterozygous mouse. Reduced tyrosine hydroxylase (TH; a marker of olfactory activity) staining on the closed side indicates completeness of naris closure. (D–F) Effects of odor exposure on H2BE level in olfactory neurons. Representative images (D) and quantification (E, left) of FLAG-H2BE in Olfr2+ neurons (D, arrowheads) exposed to odors or mineral oil (no odor). (E, right) Quantification of FLAG-H2BE in Olfr653+ neurons (chosen randomly as a negative control) exposed to odors or mineral oil (no odor). (F) Average H2BE level within neurons expressing odor-stimulated or control ORs (n = 20–60). (G,H) Representative image (G; boxed region magnified, right) and quantification (H) of the relationship between GAP43-mCherry and tyrosine hydroxylase intensities within glomeruli of an H2be-KO heterozygous mouse. Red line, best fit. *p

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Figure 2—figure supplement 1. PN responses to vinegar in flies with DTKR knockdown. ; Two-photon imaging of PN dendritic calcium responses. (A) Peak ΔF/F in glomeruli that are activated by 80% SV cider vinegar in control and DTKR knockdown flies. (B) Peak ΔF/F in the DM1 glomerulus in response to 0.2% SV cider vinegar in control and DTKR knockdown flies. (C) Representative images of calcium responses to 0.4% SV of ethyl butyrate (1:100 dilution in mineral oil). Grey-scale images show the glomerular map on the DM5 plane whereas the pseudocolored images show the change of GCaMP fluorescence. (D) Peak ΔF/F in the DM5 glomerulus in response to ethyl butyrate in control and DTKR knockdown flies. All flies have GH146-LexA, LexAop-GCaMP, Orco-Gal4, and DTKRi flies have UAS-DTKR-RNAi in addition. For each condition, n = 4–9. Error bars show s.e.m. *p < 0.05, **p < 0.01; Student's t-test comparing the response between knockdown and control groups (A) and comparing between starved and fed responses (B, D).

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