From b59c565524f864ba6aeeff2fc2148ecafdb50ab2 Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 20:46:11 +0200 Subject: [PATCH 01/11] Style gh-pages docs to match BioFSharp --- build/DocumentationTasks.fs | 4 ++-- docs/_head.html | 3 ++- docs/img/favicon.ico | Bin 0 -> 1887 bytes docs/index.md | 35 +++++++++++++++++++++++++++++++++-- 4 files changed, 37 insertions(+), 5 deletions(-) create mode 100644 docs/img/favicon.ico diff --git a/build/DocumentationTasks.fs b/build/DocumentationTasks.fs index 67e702b..85b35b8 100644 --- a/build/DocumentationTasks.fs +++ b/build/DocumentationTasks.fs @@ -12,7 +12,7 @@ let buildDocs = runDotNet (sprintf - "fsdocs build --clean --properties Configuration=Release --parameters fsdocs-package-version %s" + "fsdocs build --clean --properties Configuration=Release --parameters root https://biofsharp.com/BioFSharp.Mz/ fsdocs-package-version %s" stableVersionTag) "./" } @@ -23,7 +23,7 @@ let buildDocsPrerelease = runDotNet (sprintf - "fsdocs build --clean --properties Configuration=Release --parameters fsdocs-package-version %s" + "fsdocs build --clean --properties Configuration=Release --parameters root https://biofsharp.com/BioFSharp.Mz/ fsdocs-package-version %s" prereleaseTag) "./" } diff --git a/docs/_head.html b/docs/_head.html index 1d69afb..c6442ba 100644 --- a/docs/_head.html +++ b/docs/_head.html @@ -1 +1,2 @@ - \ No newline at end of file + + \ No newline at end of file diff --git a/docs/img/favicon.ico b/docs/img/favicon.ico new file mode 100644 index 0000000000000000000000000000000000000000..279baaa6135f15ecf491d105e450ce369bee8dbf GIT binary patch literal 1887 zcmV-l2cY;JqzqhT>+sZdoXJfs?? zEj%VXv1}VIZwWW6nZ%;-iEzl23ku)3da}&VE~m@<>9VQBqTmS#2h%?|`GeJrsSC@? zY1gv__4_{VOMI!^bX@i}nXOpF7$riGafXS~PY;8#_R~eLIMgHF{aqg})lZR^T2)qq z6-<^$lffn=YC*cEWYiuq$6xo}!;i$pA!*~T zp@AK%r=fm(x)DhqI?Y(b*<2UyIKWeLb;2X_#N4sp_m%BaDE5qgSp*{ZH zO%FJ?{lMGMHkc?(>RSnFXa7J6gQ8-8V?IAN_NAg(v?xc`wr(Dr`sta0fe z&<3>Dj^AMhpZH9r{W}t+&Z*g%EmmuLwU|aTVkA$D8p$7OL1igDJ(xLhxo&}K!~z3- zz#BlYnrpQi_+mMV(vopUifxtlWrb$t0xm5XHB$WBxVB3tf&RhFiRs$u( zDat$gu{+x^M$a%WO2myh1d?vcPj$hjWuDhuaVl6;gvFGaCsqkl!6qXM0L{Ro5l{5x zL}7CNx&pOu-PwUiCb4=WSX(xtg8_{9#;9PClSMMJNVnxDBMTf~f>)y+iPg@5WpDs7 zGlR8ZJ^Ho=?CvnSJ%sV@Y0&zb#O;=!gvwDA=CXQgNc=&q3D4y)j{g;z_yF^sZ6GCf zxD&gp1IIs4gKd+OMXrbn{pAQ3)s=cBUIUJ*HQ_J6f*pxqymuP){`7w(I@__M>v5bK zy=e&z(|J+;(#g5kOKL5+xoanwCh|fYnM^O2XbWNW-i~o<1YucqOY~PFO%l=kPI>HA z6$=b>Xn4o^+E;=)+_gBfT*mR@QOrkoqt~?{l1b!+IKZMrYIacq(4iLWo^Bk2LjZM2 zxV1slYTn%7Z967`839M+&@fm_E7WwbK3q?FX{|)NQ#0tobI`LVvliRd|^Ty>D6u^?i8zM936sdGu$CIB{WcPoT3HIL`e^QidLb>U0#`#D*BP$6WOX{xNs# z!xfiNEHEH|pV%9Eo-rTW14%P{aZNbz&B4qG6JVErXGe?e z?Y}C@AEZ?Nt!jSN+i*$ZIeSCTkIl#SAjifj?TDcr4!j3{h5RsiQ}7$bs&~QC=Xg_O z=7dP4KaZ~oE9k@X%frZI`eTDbeg6lZ#qB Date: Mon, 31 Aug 2026 20:57:34 +0200 Subject: [PATCH 02/11] Add peaks and peak arrays documentation --- docs/01_01_peaks_and_peak_arrays.fsx | 338 +++++++++ docs/data/ms2Example.mgf | 981 +++++++++++++++++++++++++++ 2 files changed, 1319 insertions(+) create mode 100644 docs/01_01_peaks_and_peak_arrays.fsx create mode 100644 docs/data/ms2Example.mgf diff --git a/docs/01_01_peaks_and_peak_arrays.fsx b/docs/01_01_peaks_and_peak_arrays.fsx new file mode 100644 index 0000000..066a97f --- /dev/null +++ b/docs/01_01_peaks_and_peak_arrays.fsx @@ -0,0 +1,338 @@ +(** +--- +title: Peaks and peak arrays +category: Spectrum processing +categoryindex: 1 +index: 1 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Peaks and peak arrays + +A mass spectrometer separates ions by their mass-to-charge ratio (m/z) and records how +many of them arrive at the detector. A peak is the recorded signal of one ion species. +In raw profile data that signal is a bump spanning many measured points around the +ion's true m/z. Centroiding reduces each such bump to a single representative pair of +an m/z value and an intensity, a step covered on the +[signal detection]({{root}}01_02_signal_detection.html) page. The `Peak` type in +BioFSharp.Mz models exactly this centroided form, and it is the smallest unit +everything else is built from. Fragment matching, database search, spectrum scoring and +similarity comparisons all consume or produce peaks. + +This page loads a measured MS2 spectrum from an mgf file, represents it as a `PeakArray`, +introduces tagged peaks and peak families (the vocabulary of fragment ion matching), and +finally converts the spectrum into the binned and sparse forms that spectrum scoring +works on. + +## Reading a spectrum from an mgf file + +Mascot generic format (mgf) is a plain text format for MS2 data. Every entry holds a +peak list plus metadata such as the precursor m/z (the m/z of the intact peptide ion +that was selected for fragmentation) and its charge state. BioFSharp ships a reader in +`BioFSharp.IO.MGF`, and the entry type with its metadata accessors lives in +`BioFSharp.FileFormats.MGF`. The example file contains a single MS2 scan of the doubly +charged peptide ANLGMEVMHER. +*) + +open BioFSharp.FileFormats.MGF +open BioFSharp.IO +open BioFSharp.Mz + +let ms2 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms2Example.mgf") + |> List.head + +printfn "%s" (MGFEntry.tryGetTitle ms2 |> Option.defaultValue "no title") + +match MGFEntry.tryGetPrecursorMZ ms2, MGFEntry.tryGetPrecursorCharges ms2 with +| Some mz, Some charges -> printfn "precursor m/z %f at charge %A" mz charges +| _ -> printfn "no precursor information" + +(** +```text +MS/MS scan at 20.93388333 min with Intensity: 501.0 and Sequence ANLGMEVMHER +precursor m/z 643.803548 at charge [2] +``` + +The reader hands back the m/z and intensity values as two separate arrays. `PeakArray.zip` +pairs them up into one `PeakArray`, an array of `Peak` values. `Peak` is a small struct +carrying `Mz` and `Intensity`, and it implements the `IPeak` interface that all peak-like +types in the library share. +*) + +let spectrum : PeakArray = PeakArray.zip ms2.Mass ms2.Intensity + +printfn "peak count: %i" spectrum.Length + +spectrum +|> Array.truncate 3 +|> Array.iter (fun p -> printfn "m/z %f intensity %f" p.Mz p.Intensity) + +(** +```text +peak count: 971 +m/z 100.666675 intensity 10.000000 +m/z 103.057515 intensity 12.000000 +m/z 103.068913 intensity 12.000000 +``` + +Some of the 971 entries are adjacent samples of the same ion signal, so the list is denser than a fully centroided spectrum, a distinction the +[signal detection]({{root}}01_02_signal_detection.html) page develops. The low +intensity values are typical for a single MS2 scan of a low abundance precursor. + +## Creating and transforming peaks + +Single peaks are created with `Peaks.createPeak`. Because `PeakArray` is a plain array +under the hood, the familiar `Array` functions work on it, and `PeakArray.map` builds a +new peak array from an existing one. A common use is normalizing all intensities to the +most intense peak of the spectrum (the base peak), which makes spectra of different +absolute intensity comparable. +*) + +let singlePeak = Peaks.createPeak 175.119 24. +printfn "m/z %f intensity %f" singlePeak.Mz singlePeak.Intensity + +let basePeakIntensity = + spectrum |> Array.map (fun p -> p.Intensity) |> Array.max + +let normalized = + spectrum + |> PeakArray.map (fun p -> Peaks.createPeak p.Mz (p.Intensity / basePeakIntensity)) + +normalized +|> Array.truncate 3 +|> Array.iter (fun p -> printfn "m/z %f relative intensity %f" p.Mz p.Intensity) + +(** +```text +m/z 175.119000 intensity 24.000000 +m/z 100.666675 relative intensity 0.058824 +m/z 103.057515 relative intensity 0.070588 +m/z 103.068913 relative intensity 0.070588 +``` + +`PeakList` is the list-backed sibling of `PeakArray` with the same `zip` and `map` +functions. When you need the raw m/z and intensity sequences back, for example +to feed a plotting library, `PeakList.unzip` splits a peak list into its two component +lists. +*) + +let mzList, intensityList = + spectrum |> List.ofArray |> PeakList.unzip + +printfn "m/z values: %A" (mzList |> List.truncate 3) +printfn "intensity values: %A" (intensityList |> List.truncate 3) + +(** +```text +m/z values: [100.666675; 103.057515; 103.068913] +intensity values: [10.0; 12.0; 12.0] +``` + +## Tagging peaks with ion types + +When a peptide is fragmented in the mass spectrometer, its backbone breaks at +characteristic positions. A fragment that keeps the N-terminus is called a b ion, one +that keeps the C-terminus a y ion (there are more series, a, c, x and z). A peak on its +own does not know which fragment it belongs to. For fragment matching the library +attaches that information as a tag: `Ions.IonTypeFlag` is a flags enum naming the ion +series, `TaggedMass` pairs a flag with a mass, and `TaggedPeak.TaggedPeak` pairs a flag +with an m/z and an intensity. + +The tags become concrete with two fragments of ANLGMEVMHER, computed by hand from the +monoisotopic residue masses. The b2 ion covers the first two residues, the y1 ion is the +C-terminal arginine. +*) + +// b2 of ANLGMEVMHER: A + N + proton = 71.03711 + 114.04293 + 1.00728 +let b2 = TaggedMass.createTaggedMass Ions.IonTypeFlag.B 186.08732 + +// y1 of ANLGMEVMHER: R + H2O + proton = 156.10111 + 18.01056 + 1.00728 +let y1 = TaggedMass.createTaggedMass Ions.IonTypeFlag.Y 175.11895 + +printfn "%A ion at m/z %f" b2.Iontype b2.Mass +printfn "%A ion at m/z %f" y1.Iontype y1.Mass + +(** +```text +B ion at m/z 186.087320 +Y ion at m/z 175.118950 +``` + +Fragments often show up together with satellite peaks, for example the same ion minus a +water molecule (18.01056 Da lighter). `TaggedMass.createTaggedH2OLoss` builds such a +loss tag by combining the ion series flag with the `lossH2O` flag, and +`Peaks.createPeakFamily` groups a main peak with its dependent satellite peaks into a +`PeakFamily`. The [in silico fragmentation]({{root}}02_01_fragmentation.html) page +produces exactly this shape, `PeakFamily` values for every fragment of a +peptide, so the type is introduced here on a small example. +*) + +let y1WaterLoss = + TaggedMass.createTaggedH2OLoss Ions.IonTypeFlag.Y (175.11895 - 18.01056) + +let y1Family = Peaks.createPeakFamily y1 [ y1WaterLoss ] + +printfn "main peak: %A at m/z %f" y1Family.MainPeak.Iontype y1Family.MainPeak.Mass +y1Family.DependentPeaks +|> List.iter (fun t -> printfn "dependent peak: %A at m/z %f" t.Iontype t.Mass) + +(** +```text +main peak: Y at m/z 175.118950 +dependent peak: Y, lossH2O at m/z 157.108390 +``` + +The dependent peak carries both flags at once, which is what a flags enum is for. A +matching function can later ask for the series with `Ions.hasFlag`. + +Because ANLGMEVMHER is the peptide that was actually fragmented in our example scan, its +y1 ion should be present in the measured spectrum. It is. +*) + +spectrum +|> Array.filter (fun p -> abs (p.Mz - y1.Mass) < 0.05) +|> Array.iter (fun p -> printfn "measured peak near y1: m/z %f intensity %f" p.Mz p.Intensity) + +(** +```text +measured peak near y1: m/z 175.118252 intensity 12.000000 +measured peak near y1: m/z 175.120109 intensity 12.000000 +measured peak near y1: m/z 175.121966 intensity 24.000000 +``` + +A `TaggedPeak.TaggedPeak` adds an intensity to the tag and implements `IPeak` like every +other peak type, so tagged peaks fit into a `PeakArray` as well. +*) + +let y1Peak = TaggedPeak.createTaggedPeak Ions.IonTypeFlag.Y 175.11895 24. +printfn "%A peak at m/z %f with intensity %f" y1Peak.Iontype y1Peak.Mz y1Peak.Intensity + +(** +```text +Y peak at m/z 175.118950 with intensity 24.000000 +``` + +## Binning a spectrum into unit dalton bins + +SEQUEST-style scoring compares a measured spectrum against a predicted one as vectors. +For that the continuous m/z axis is discretized into bins of 1 Da width, and every peak +is assigned to its nearest bin. `PeakArray.peaksToNearestUnitDaltonBinVector` performs +this conversion between a lower and an upper mass border and returns an `FSharp.Stats` +vector. When several peaks fall into the same bin, the bin keeps the highest intensity. +The produced vector has length `maxMassBoarder - minMassBoarder`, here 900 bins covering +m/z 100 to 1000. +*) + +open FSharp.Stats + +let binned = PeakArray.peaksToNearestUnitDaltonBinVector spectrum 100.0 1000.0 + +printfn "vector length: %i" (Vector.length binned) + +let occupied = + binned + |> Vector.toArray + |> Array.indexed + |> Array.filter (fun (_, intensity) -> intensity > 0.) + +printfn "occupied bins: %i" occupied.Length + +occupied +|> Array.truncate 5 +|> Array.iter (fun (i, intensity) -> printfn "bin %i (m/z %i): %f" i (i + 100) intensity) + +(** +```text +vector length: 900 +occupied bins: 378 +bin 1 (m/z 101): 10.000000 +bin 3 (m/z 103): 12.000000 +bin 4 (m/z 104): 12.000000 +bin 10 (m/z 110): 24.000000 +bin 12 (m/z 112): 10.000000 +``` + +971 peaks collapse into 378 occupied bins because peaks closer together than 1 Da share +a bin. The bin index is simply the rounded m/z minus the lower mass border, so bin 0 +collects the peaks around m/z 100. + +## Comparing spectra with sparse peak arrays + +Most of the 900 bins above are zero. `SparsePeakArray` stores only the occupied bins in +a dictionary from bin index to intensity, which saves memory and makes comparing two +spectra a walk over one dictionary with lookups in the other. +`SparsePeakArray.peaksToNearestBinVector` builds one from a peak array. The first two +arguments control the binning: with a bin width of 1.0 and an offset of 0.5 every peak +lands in its nearest unit bin. In the sparse form, peaks sharing a bin are summed. The +record also carries the two conversion functions `MzToBinIdx` and `BinIdxToMz` so you +can move between m/z values and bin indices. +*) + +let sparse = + spectrum + |> SparsePeakArray.peaksToNearestBinVector 1.0 0.5 100.0 1000.0 + +printfn "occupied bins: %i" sparse.Data.Count +printfn "bin index of the y1 m/z: %i" (sparse.MzToBinIdx 175.11895) + +(** +```text +occupied bins: 378 +bin index of the y1 m/z: 175 +``` + +`SparsePeakArray.dot` computes the dot product of two sparse spectra: for every bin both +spectra occupy, it multiplies the intensities and sums the products. The dot product +(usually after normalization) is the standard similarity measure between binned spectra, +and a spectrum compared with itself gives the maximal value. + +The example file contains only one scan, so we simulate a spectrum of an unrelated +peptide by shifting every measured m/z by 7.3 Da. The shifted copy has the same +intensity distribution, only in different bins. +*) + +// same peaks moved by 7.3 Da: a stand-in for a spectrum of an unrelated peptide +let shifted = + spectrum + |> PeakArray.map (fun p -> Peaks.createPeak (p.Mz + 7.3) p.Intensity) + +let sparseShifted = + shifted + |> SparsePeakArray.peaksToNearestBinVector 1.0 0.5 100.0 1000.0 + +printfn "self dot product: %f" (SparsePeakArray.dot sparse sparse) +printfn "cross dot product: %f" (SparsePeakArray.dot sparse sparseShifted) + +(** +```text +self dot product: 3692147.000000 +cross dot product: 163836.000000 +``` + +The spectrum agrees with itself far better than with the shifted copy, exactly what a +similarity measure should say. The remaining cross product comes from bins where a +shifted peak happens to land on another real peak. + +The next step is generating predicted fragment peaks to match against, the topic of the +[in silico fragmentation]({{root}}02_01_fragmentation.html) page. +*) diff --git a/docs/data/ms2Example.mgf b/docs/data/ms2Example.mgf new file mode 100644 index 0000000..48fa908 --- /dev/null +++ b/docs/data/ms2Example.mgf @@ -0,0 +1,981 @@ +BEGIN IONS +CHARGE=2+ +PEPMASS=1285.591258 +PRECURSORMZ=643.8035478 +TITLE=MS/MS scan at 20.93388333 min with Intensity: 501.0 and Sequence ANLGMEVMHER + + +100.666675 10.000000 +103.057515 12.000000 +103.068913 12.000000 +104.052913 12.000000 +104.054345 12.000000 +104.058640 12.000000 +110.065394 12.000000 +110.072756 12.000000 +110.075700 24.000000 +110.080118 12.000000 +112.081325 10.000000 +113.067050 12.000000 +113.073019 12.000000 +113.074511 24.000000 +114.882914 10.000000 +115.045425 10.000000 +117.069808 10.000000 +117.311367 10.000000 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+1129.490806 12.000000 +1130.604176 10.000000 +1187.552274 10.000000 +1187.890833 10.000000 +1198.526706 10.000000 +1215.528204 10.000000 +1226.527637 12.000000 +1226.552212 12.000000 +1226.576787 12.000000 +1337.629577 10.000000 + + +END IONS From c8b601b5207cdd30dcba289f508b975115b3365f Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 21:12:53 +0200 Subject: [PATCH 03/11] Add signal detection documentation --- docs/01_02_signal_detection.fsx | 274 + docs/data/ms1Example.mgf | 78177 ++++++++++++++++++++++++++++++ 2 files changed, 78451 insertions(+) create mode 100644 docs/01_02_signal_detection.fsx create mode 100644 docs/data/ms1Example.mgf diff --git a/docs/01_02_signal_detection.fsx b/docs/01_02_signal_detection.fsx new file mode 100644 index 0000000..5028aa7 --- /dev/null +++ b/docs/01_02_signal_detection.fsx @@ -0,0 +1,274 @@ +(** +--- +title: Signal detection and centroiding +category: Spectrum processing +categoryindex: 1 +index: 2 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Signal detection and centroiding + +A mass spectrometer in profile mode writes down the detector signal as a +continuous-looking trace: many m/z and intensity pairs per true ion signal. Every ion +species shows up as a roughly Gaussian bump that is a few dozen points wide, with its +apex at the ion's m/z. Charge state determination, database search, spectrum scoring and +quantification all expect one peak per ion species. Centroiding reduces every bump to a +single representative peak, an m/z position plus an apex or summed intensity. + +`SignalDetection` implements a wavelet-based peak picker modeled after French et al. +(doi: [10.1021/pr500886y](https://doi.org/10.1021/pr500886y)). The idea: a Mexican hat +wavelet (a Ricker wavelet, which looks like a Gaussian bump with dips on both sides) is +slid along the m/z axis at several widths and correlated with the intensity trace. +Wherever the trace contains a bump of matching width, the correlation is high. True +peaks score well across scales while single-point noise spikes do not, which is what +makes the approach less trigger-happy than simple local maxima. This page covers the +wavelet approach. + +## Loading a profile MS1 spectrum + +The example file holds one MS1 survey scan (a full scan over the precursor mass range, +recorded before the instrument picks ions for fragmentation) stored in mgf format. We +load it with the reader from BioFSharp, as on the +[peaks and peak arrays]({{root}}01_01_peaks_and_peak_arrays.html) page, and check +empirically that it really is profile data by looking at the number of points and the +spacing between neighboring points. +*) + +open BioFSharp.FileFormats.MGF +open BioFSharp.IO +open BioFSharp.Mz + +let ms1 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms1Example.mgf") + |> List.head + +let profileMz = ms1.Mass +let profileIntensity = ms1.Intensity + +printfn "%s" (MGFEntry.tryGetTitle ms1 |> Option.defaultValue "no title") +printfn "points in scan: %i" profileMz.Length +printfn "m/z range: %f to %f" profileMz.[0] (Array.last profileMz) + +let spacings = + profileMz |> Array.pairwise |> Array.map (fun (a, b) -> b - a) + +spacings +|> Array.skip 2 +|> Array.truncate 3 +|> Array.iter (printfn "spacing between adjacent points: %f") + +printfn "median spacing: %f" ((Array.sort spacings).[spacings.Length / 2]) +printfn "largest gap: %f" (Array.max spacings) + +(** +```text +MS scan at 20.91351667 min +points in scan: 78170 +m/z range: 400.001711 to 1249.609952 +spacing between adjacent points: 0.005613 +spacing between adjacent points: 0.005614 +spacing between adjacent points: 0.002807 +median spacing: 0.003504 +largest gap: 1.481318 +``` + +78170 points with a median spacing of about 0.0035 Da is profile data. A centroided MS1 +scan of the same range would have a few hundred to a few thousand peaks, one per +detected ion signal. The largest gap tells the second half of the story: the +instrument leaves out stretches where it saw nothing, so the sampling is dense inside +peak regions and sparse between them. + +## Padding sparse regions + +The wavelet correlation assumes reasonably dense, gap-free sampling, because the width +of the wavelet is derived from the local point spacing. A 1.5 Da hole in the data would +inflate that estimate and distort the correlation around the hole. `Padding.paddDataBy` +therefore fills sparse regions with artificial points at a fixed base intensity before +the peak picker runs. + +Its parameters, built with `Padding.createPaddingParameters`: `paddingYValue` is the +intensity written into every inserted point, typically 0. or the minimum intensity of +the data. `maximumPaddingPoints` caps how many points may be inserted into one gap, +and `None` fills gaps completely. `mzTolerance` sets the m/z distance at which the +inserted points are placed, here 0.05 Da, finer than any real peak width. +`windowSize` and `spacingPerc` set the points per window and the percentile of the +spacings inside it used to estimate the typical local point spacing. A spacing is +padded as a gap when it stands out against this local estimate and exceeds +`mzTolerance`. +*) + +let paddingParams = + SignalDetection.Padding.createPaddingParameters 0. None 0.05 150 95. + +let paddedMz, paddedIntensity = + SignalDetection.Padding.paddDataBy paddingParams profileMz profileIntensity + +printfn "points before padding: %i" profileMz.Length +printfn "points after padding: %i" paddedMz.Length + +(** +```text +points before padding: 78170 +points after padding: 83672 +``` + +The added points all carry intensity 0. and only exist so the wavelet sees a continuous +baseline between the real peak regions. + +## Centroiding with the wavelet peak picker + +`Wavelet.toCentroidWithRicker2D` takes a `WaveletParameters` record and the padded +arrays and returns the centroided m/z and intensity arrays. The parameters, built with +`Wavelet.createWaveletParameters`: + +`numberOfScales` is the number of wavelet widths tried at every position. The widths +span a fixed range from one to seven times the local point spacing, so larger values +sample that range more finely at more cost. `yThreshold` is an absolute intensity +floor below which a bump's apex is dropped, and with 0. the filtering is left to the +signal-to-noise criterion. `mzTolerance` is the grouping tolerance in m/z: candidate +peaks closer together than this compete, and only the best-scoring one survives. +`sNRS_Percentile` sets which percentile of the local correlation scores is taken as +the noise estimate, and `minSNR` is the minimum ratio between a candidate's +correlation score and that estimate. `refineMZ` reports the m/z as the +intensity-weighted mean over the bump (`true`) or the position of the apex point +(`false`), and `sumIntensities` reports the intensity as the sum over the bump +(`true`) or the apex intensity (`false`). + +The values below are what a production proteomics pipeline uses for MS1 scans: 3 +scales, mzTolerance 0.1, the 95th percentile for the noise estimate and a minimum SNR +of 1. For the denser features of MS2 spectra the same pipeline raises the number of +scales to 10. We report apex positions and apex intensities first. +*) + +let waveletParams = + SignalDetection.Wavelet.createWaveletParameters 3 0. 0.1 95. 1. false false + +let centroidMz, centroidIntensity = + SignalDetection.Wavelet.toCentroidWithRicker2D waveletParams paddedMz paddedIntensity + +printfn "profile points: %i" profileMz.Length +printfn "centroids: %i" centroidMz.Length + +(** +```text +profile points: 78170 +centroids: 1454 +``` + +78170 profile points become 1454 centroids, one per detected ion signal, a reduction of +roughly a factor of fifty on this scan. + +## Watching a bump collapse to a centroid + +Numbers over the whole scan hide what happens locally, so we zoom into a 1.5 Da window +around the most intense signal of the scan. First the profile points near its apex. +*) + +let inWindow lo hi (mz, _) = mz >= lo && mz <= hi + +let profileWindow = + Array.zip profileMz profileIntensity + |> Array.filter (inWindow 434.9 436.4) + +printfn "profile points between m/z 434.9 and 436.4: %i" profileWindow.Length + +profileWindow +|> Array.filter (inWindow 435.18 435.21) +|> Array.iter (fun (mz, i) -> printfn "m/z %f intensity %8.1f" mz i) + +(** +```text +profile points between m/z 434.9 and 436.4: 398 +m/z 435.180270 intensity 1791.0 +m/z 435.183197 intensity 4332.0 +m/z 435.186125 intensity 7143.0 +m/z 435.189052 intensity 10025.0 +m/z 435.191980 intensity 9749.0 +m/z 435.194907 intensity 7993.0 +m/z 435.197835 intensity 6344.0 +m/z 435.200762 intensity 3923.0 +m/z 435.203690 intensity 2902.0 +m/z 435.206617 intensity 1836.0 +m/z 435.209545 intensity 1337.0 +``` + +The intensity rises point by point to an apex at m/z 435.189 and falls off again, the +Gaussian bump from the introduction in the flesh. Now the centroids the wavelet produced +for the same window. +*) + +Array.zip centroidMz centroidIntensity +|> Array.filter (inWindow 434.9 436.4) +|> Array.iter (fun (mz, i) -> printfn "m/z %f intensity %8.1f" mz i) + +(** +```text +m/z 435.189052 intensity 10025.0 +m/z 435.525777 intensity 5519.0 +m/z 435.856773 intensity 2806.0 +m/z 436.190825 intensity 1053.0 +``` + +398 profile points collapse to four centroids. The first sits exactly on the apex we +just looked at. The four peaks are spaced about 0.334 Da apart with intensities falling +off from left to right, which is the signature of an isotope cluster. Each following +peak is the same ion species with one more heavy isotope, and since the mass +difference between carbon-13 and carbon-12 is about 1.0034 Da, a spacing of +1.0034 / 3 means the ion carries three charges. + +Rerunning with `refineMZ` and `sumIntensities` turned on changes what each centroid +reports. The m/z becomes the intensity-weighted mean over the bump and the intensity +becomes the sum over all its profile points, which is the better quantity when peak +areas are compared later. +*) + +let summingParams = + SignalDetection.Wavelet.createWaveletParameters 3 0. 0.1 95. 1. true true + +let refinedMz, refinedIntensity = + SignalDetection.Wavelet.toCentroidWithRicker2D summingParams paddedMz paddedIntensity + +Array.zip refinedMz refinedIntensity +|> Array.filter (inWindow 434.9 436.4) +|> Array.iter (fun (mz, i) -> printfn "m/z %f intensity %8.1f" mz i) + +(** +```text +m/z 435.192899 intensity 59306.0 +m/z 435.526224 intensity 31209.0 +m/z 435.859807 intensity 17631.0 +m/z 436.193927 intensity 6746.0 +``` + +Same four signals, now with summed intensities and slightly shifted m/z positions +because the weighted mean also feels the asymmetric tails of each bump. + +## Where the centroids go next + +The centroided arrays can be zipped back into a `PeakArray` (see +[peaks and peak arrays]({{root}}01_01_peaks_and_peak_arrays.html)). The isotope spacing +we read off by eye above is what +[charge state determination]({{root}}01_03_charge_state_determination.html) exploits +systematically to assign each precursor its charge, needed to turn an m/z value into a +peptide mass for the database search. +*) diff --git a/docs/data/ms1Example.mgf b/docs/data/ms1Example.mgf new file mode 100644 index 0000000..ad754b2 --- /dev/null +++ b/docs/data/ms1Example.mgf @@ -0,0 +1,78177 @@ +BEGIN IONS +TITLE=MS scan at 20.91351667 min + + +400.001711 9.000000 +400.041004 59.000000 +400.071880 9.000000 +400.077493 9.000000 +400.083107 14.000000 +400.085914 14.000000 +400.091528 9.000000 +400.094335 24.000000 +400.097142 9.000000 +400.099949 9.000000 +400.102756 9.000000 +400.119598 19.000000 +400.122405 14.000000 +400.130826 14.000000 +400.133633 9.000000 +400.136440 9.000000 +400.139247 9.000000 +400.142055 47.000000 +400.144862 14.000000 +400.147669 14.000000 +400.153283 14.000000 +400.156090 23.000000 +400.158897 23.000000 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9.000000 + + +END IONS From 9ca89d48313d038c690d3aaca6ea811bae317e6f Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 21:25:56 +0200 Subject: [PATCH 04/11] Add charge state determination documentation --- docs/01_03_charge_state_determination.fsx | 340 ++++++++++++++++++++++ 1 file changed, 340 insertions(+) create mode 100644 docs/01_03_charge_state_determination.fsx diff --git a/docs/01_03_charge_state_determination.fsx b/docs/01_03_charge_state_determination.fsx new file mode 100644 index 0000000..6cf5b11 --- /dev/null +++ b/docs/01_03_charge_state_determination.fsx @@ -0,0 +1,340 @@ +(** +--- +title: Charge state determination +category: Spectrum processing +categoryindex: 1 +index: 3 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Charge state determination + +A mass spectrometer reports m/z, the mass of an ion divided by the number of charges +it carries. The database search at the end of the pipeline works with neutral peptide +masses, so at some point the charge z has to be inferred. The stakes are concrete: a precursor observed at m/z 643.8 weighs +about 1285.6 Da if it carries two charges and about 1928.4 Da if it carries three. +Search the wrong mass range and the peptide is never found. + +The information needed to infer z is already in the MS1 scan. Elements come in +naturally occurring isotopes, and with roughly one in a hundred carbon atoms being a +heavy ¹³C, a population of identical peptide molecules spreads out into an isotope +cluster: the monoisotopic peak, a peak one neutron heavier, one two neutrons heavier, +and so on. For a peptide with neutral mass M at charge z, the n-th isotope peak sits +at m/z = (M + z·H + n·1.00335) / z. Adjacent cluster peaks are therefore spaced +1.00335 / z apart on the m/z axis. A cluster whose peaks sit 1.0 apart is singly +charged, 0.5 apart means 2+, 0.33 apart means 3+, 0.25 apart means 4+. The +`ChargeState` module reads the +charge out of exactly this spacing, using 1 / z as an approximation of the +theoretical distance. + +`ms2Example.mgf` holds a fragment +spectrum of the peptide ANLGMEVMHER, recorded at 20.93 min. `ms1Example.mgf` is the +MS1 survey scan recorded right before it, at 20.91 min in the same run, where the +fragmented precursor was measured intact, so its isotope cluster is sitting in that +scan's data. We determine the charge from the MS1 scan and check the resulting mass +against what the MS2 file states in its header. +*) + +open BioFSharp.FileFormats.MGF +open BioFSharp.IO +open BioFSharp.Mz + +let ms2 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms2Example.mgf") + |> List.head + +let precursorMZ = + match MGFEntry.tryGetPrecursorMZ ms2 with + | Some mz -> mz + | None -> failwith "no precursor m/z in the MS2 header" + +printfn "%s" (MGFEntry.tryGetTitle ms2 |> Option.defaultValue "no title") +printfn "precursor m/z: %f" precursorMZ + +(** +```text +MS/MS scan at 20.93388333 min with Intensity: 501.0 and Sequence ANLGMEVMHER +precursor m/z: 643.803548 +``` + +The header also carries a `PEPMASS` of 1285.591258 Da. That value is our ground +truth. If the charge determination works, we arrive at this mass using only the MS1 +peaks and the precursor m/z. + +## Centroiding the MS1 survey scan + +Charge determination runs on centroided peaks, one peak per ion signal. We prepare +the MS1 scan exactly as on the +[signal detection]({{root}}01_02_signal_detection.html) page, with the same padding +and wavelet parameters, and refer there for what each of them does. +*) + +let ms1 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms1Example.mgf") + |> List.head + +let paddingParams = + SignalDetection.Padding.createPaddingParameters 0. None 0.05 150 95. + +let waveletParams = + SignalDetection.Wavelet.createWaveletParameters 3 0. 0.1 95. 1. false false + +let centroidMz, centroidIntensity = + SignalDetection.Padding.paddDataBy paddingParams ms1.Mass ms1.Intensity + ||> SignalDetection.Wavelet.toCentroidWithRicker2D waveletParams + +printfn "centroids: %i" centroidMz.Length + +(** +```text +centroids: 1454 +``` + +Before handing these arrays to the module, we look at the centroids in a small +window starting at the precursor m/z, together with their spacings. +*) + +let clusterWindow = + Array.zip centroidMz centroidIntensity + |> Array.filter (fun (mz, _) -> mz >= 643.5 && mz <= 645.5) + +clusterWindow +|> Array.iter (fun (mz, i) -> printfn "m/z %f intensity %5.1f" mz i) + +clusterWindow +|> Array.map fst +|> Array.pairwise +|> Array.iter (fun (a, b) -> printfn "spacing: %f" (b - a)) + +(** +```text +m/z 643.802071 intensity 501.0 +m/z 644.311347 intensity 352.0 +m/z 644.810133 intensity 150.0 +m/z 645.305547 intensity 124.0 +spacing: 0.509276 +spacing: 0.498786 +spacing: 0.495414 +``` + +A falling intensity ladder spaced at about half a dalton, which by the formula above +is the signature of a doubly charged ion. The first peak's intensity of 501.0 is the +precursor intensity the MS2 title records, so we are looking at the right signal. +Reading the charge off by eye works here. The module does the same thing +systematically and with safeguards for messier neighborhoods. + +## Generating charge state candidates + +`ChargeState.putativePrecursorChargeStatesBy` needs a `ChargeDetermParams` record, +built with `createChargeDetermParams`. Its parameters, in order: the expected minimal +and maximal charge span the states that are considered at all. Tryptic peptides +mostly ionize at 2+ to 4+, one proton at the N-terminus and one at the C-terminal +lysine or arginine, sometimes more on internal basic residues, so 2 and 4 are +reasonable bounds. `Width` is the m/z window, in daltons, scanned to the right of the +precursor peak for cluster members. 1.1 covers one full isotope spacing at charge 1 +and correspondingly more peaks at higher charges. The next two values decide which +peaks in that window count as cluster members: a peak must reach `MinIntensity` +(here 15 percent) of the start peak's intensity and `DeltaMinIntensity` (here 30 +percent) of the peak before it, which cuts the cluster off once it has decayed into +noise. `NrOfRndSpectra` is the number of random clusters simulated for the hypothesis +test in the last section. The module author's design uses 10000. +*) + +let chargeParams = + ChargeState.createChargeDetermParams 2 4 1.1 0.15 0.3 10000 + +(** +The function locates the centroid closest to the given precursor m/z, collects the +window peaks that pass the two intensity criteria, and then considers every subset of +them that contains the start peak, because chemical noise can sit between real +isotope peaks and a fixed left-to-right reading would stumble over it. Each subset +gets the charge from the allowed range whose theoretical spacing best matches the +subset's mean peak spacing, a deviation measure (`MZChargeDev`) quantifying how well +the spacings fit that charge, and a score, the weighted sum of that deviation and a +length quotient, the fraction of window peaks the subset uses (weights from a linear +discriminant analysis, lower is better). The neutral mass `PutMass` is computed from the precursor m/z and the +assigned charge with `Mass.ofMZ` from BioFSharp. The result keeps the best subset per +distinct charge, sorted by score. One safeguard to know about: a window holding 15 or +more peaks is considered too noisy for spacing analysis, and the function then falls +back to enumerating every allowed charge instead. + +The two string arguments are free-text identifiers for the MS1 and MS2 scan. They are +carried into the result record so a downstream consumer can trace which spectra an +assignment belongs to. +*) + +let printCandidate (c: ChargeState.AssignedCharge) = + printfn "charge %i+ score %8.4f mzChargeDev %8.4f putMass %9.4f peaks %i" + c.PrecCharge c.Score c.MZChargeDev c.PutMass c.SubSetLength + +let putativeCharges = + ChargeState.putativePrecursorChargeStatesBy + chargeParams centroidMz centroidIntensity + "MS1 at 20.91 min" "MS2 at 20.93 min" precursorMZ + +putativeCharges |> List.iter printCandidate + +(** +```text +charge 2+ score -0.2906 mzChargeDev 0.0094 putMass 1285.5925 peaks 3 +``` + +One candidate survives the subset competition: charge 2, backed by a cluster of three +peaks (the fourth ladder peak from above lies beyond the 1.1 Da window). The putative +mass of 1285.5925 Da lands within about a millidalton of the 1285.591258 Da stated in +the MS2 header, the small residual coming down to which mass constant is used for the +added protons. The mass the database search needs has been recovered from the MS1 +peak pattern alone. + +## Selecting among candidates + +`ChargeState.removeSubSetsOfBestHit` takes the first list element +as the best hit, which the score-sorted output of `putativePrecursorChargeStatesBy` +provides, and drops every other candidate whose peak set is a subset of the best +hit's peaks. Such candidates explain no signal the best hit does not already explain. +Candidates built from different peaks survive, since they may describe a second, +overlapping cluster. +*) + +let assignedCharges = + ChargeState.removeSubSetsOfBestHit putativeCharges + +assignedCharges |> List.iter printCandidate + +(** +```text +charge 2+ score -0.2906 mzChargeDev 0.0094 putMass 1285.5925 peaks 3 +``` + +With a single candidate there is nothing to remove. The filter becomes visible on a +second cluster. + +## Cross-checking on the cluster at m/z 435.19 + +The [signal detection]({{root}}01_02_signal_detection.html) page ended on the most +intense signal of this scan, four centroids around m/z 435.19 spaced about 0.334 Da +apart, and read a charge of 3 off that spacing by eye. The same call reproduces this +on a second, independent case. +*) + +let putativeCharges435 = + ChargeState.putativePrecursorChargeStatesBy + chargeParams centroidMz centroidIntensity + "MS1 at 20.91 min" "-" 435.189052 + +putativeCharges435 |> List.iter printCandidate + +(** +```text +charge 3+ score -0.2950 mzChargeDev 0.0050 putMass 1302.5453 peaks 3 +charge 2+ score 0.0372 mzChargeDev 0.2372 putMass 868.3636 peaks 2 +``` + +The best candidate is the expected 3+, built from three cluster peaks. The 2+ +candidate is an artifact of the subset construction: the start peak paired with the +second isotope peak alone spans 0.668 Da, read as one poor charge-2 gap, which both +its deviation and its score reflect. Its two peaks are contained in the best hit's +three, so the subset filter disposes of it. +*) + +ChargeState.removeSubSetsOfBestHit putativeCharges435 +|> List.iter printCandidate + +(** +```text +charge 3+ score -0.2950 mzChargeDev 0.0050 putMass 1302.5453 peaks 3 +``` + +## Testing the cluster against random spacings + +`MZChargeDev` is a unitless deviation, and on its own it is hard to say how small is +small enough. The module frames the question as a hypothesis test. The null +hypothesis says the cluster is a random arrangement of legal charge spacings. When +random arrangements practically never fit the assigned charge as well as the observed +cluster does, the null is rejected and the cluster counts as a genuine isotope +pattern. First, `peakPosStdDevBy` estimates the spacing scatter of the current +measurement from the candidates' deviations between real and theoretical peak +spacings. +*) + +let peakPosStdDev = + ChargeState.peakPosStdDevBy putativeCharges + +printfn "peak position standard deviation: %.4f" peakPosStdDev + +(** +```text +peak position standard deviation: 0.0074 +``` + +`initMzDevOfRndSpec` then builds a generator of random clusters: each simulated +cluster has as many peaks as the observed one, and every gap is computed as one over +a jittered charge (the implementation draws a charge from 1 up to one below the +configured maximum and adds normal jitter with the standard deviation just estimated +to the charge before taking the reciprocal). `empiricalPValueOfSim` generates `NrOfRndSpectra` such clusters +(memoized per peak count and charge, so repeated calls are cheap) and returns the +fraction of them whose deviation from the assigned charge's spacing is at most as +large as the observed one. That fraction is the empirical p-value. The design +described in the module author's thesis tests each candidate that survives the subset +filter this way, against a distribution of 10000 random clusters, and rejects the +null hypothesis at a significance criterion of 0.05. +*) + +let rnd = System.Random() + +let mzDevOfRandomCluster = + ChargeState.initMzDevOfRndSpec rnd chargeParams peakPosStdDev + +let bestHit = assignedCharges.Head + +let pValue = + ChargeState.empiricalPValueOfSim + mzDevOfRandomCluster + (bestHit.SubSetLength, float bestHit.PrecCharge) + bestHit.MZChargeDev + +printfn "empirical p-value for charge %i+: %.2f" bestHit.PrecCharge pValue + +(** + +```text +empirical p-value for charge 2+: 0.25 +``` + +Part of the simulation draws its jitter from a random source the caller cannot seed, +so this number varies slightly between runs. It lies far above the 0.05 criterion, so +the null hypothesis stands and a pipeline applying the thesis criterion would drop +this candidate on the position metric alone. That is the expected verdict for a +cluster this short: every random gap is itself a legal charge spacing, and with only +two gaps to check, roughly a quarter of the random clusters draw the matching charge +twice and fit at least as well as the real measurement. Production pipelines +therefore also lean on the score ranking and assume typical charges such as 2+ and 3+ +as a fallback when no candidate is convincing. The computed value can be stored in +the `PositionMetricPValue` field for exactly this kind of filtering. + +## Where the charge goes next + +Charge and mass together are what the next stage consumes, whether they come from a +ranked candidate like this one or from the 2+ and 3+ fallback: the mass selects the +candidate peptides to score against the fragment spectrum, the subject of +[peptide search databases]({{root}}02_02_search_databases.html). +*) From e1ecb84a242faf538b5a3adfaf4472c588311db4 Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 21:36:53 +0200 Subject: [PATCH 05/11] Add in silico fragmentation documentation --- docs/02_01_fragmentation.fsx | 350 +++++++++++++++++++++++++++++++++++ 1 file changed, 350 insertions(+) create mode 100644 docs/02_01_fragmentation.fsx diff --git a/docs/02_01_fragmentation.fsx b/docs/02_01_fragmentation.fsx new file mode 100644 index 0000000..cfa61fa --- /dev/null +++ b/docs/02_01_fragmentation.fsx @@ -0,0 +1,350 @@ +(** +--- +title: In silico fragmentation +category: Peptide identification +categoryindex: 2 +index: 1 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# In silico fragmentation + +An MS2 spectrum is the fragment spectrum of one selected peptide ion. Inside the +collision cell the peptide collides with gas molecules until its backbone breaks, +and the instrument records the m/z values of the resulting pieces. Reading the +sequence directly out of those peaks is hard. Search engines therefore go the +other way: take a candidate sequence and predict which fragment masses it would +produce, then check how many of the predictions show up in the measured +spectrum. The candidate whose predictions explain the spectrum best wins. This +page covers the prediction side, implemented in the `Fragmentation` module. + +Under collision-induced dissociation (CID) and electron transfer dissociation (ETD) +the backbone cleaves between two residues, leaving an N-terminal piece and a +C-terminal piece. The Roepstorff-Fohlman-Biemann nomenclature names the +fragments by the exact bond that broke: N-terminal fragments are called a, b or +c ions, C-terminal fragments x, y or z ions. The index counts residues from the +fragment's own terminus, so b2 covers the first two residues and y1 is the last +residue alone. CID, the most common fragmentation method and the plausible origin +of our example spectrum, produces mostly b and y ions, which is why those two +series carry most identifications. + +On top of the backbone fragments come satellite peaks. A fragment containing +arginine, lysine, asparagine or glutamine can additionally lose ammonia (17 Da), +and a fragment containing serine, threonine, glutamate or aspartate can lose +water (18 Da). These losses appear as smaller peaks slightly below the main +fragment peak. The generator knows both rules, the residue sets live in the +module as `Fragmentation.aminoLossSet` and `Fragmentation.waterLossSet`. + +`ms2Example.mgf` holds a measured MS2 scan of the doubly charged peptide ANLGMEVMHER, and the +[charge state determination]({{root}}01_03_charge_state_determination.html) page +recovered its neutral mass. Here we predict the fragments of ANLGMEVMHER and +show that they actually sit in that spectrum. + +## Building the peptide + +The fragmentation functions work on an amino acid list from BioFSharp. +`BioList.ofAminoAcidString` parses the one letter sequence. Every `Series` +function additionally takes a mass function of type `IBioItem -> float`, which +is how the caller decides whether the ladder is computed from monoisotopic or +average masses. We pass `BioItem.monoisoMass`, since fragment matching at 0.05 +Da tolerance needs monoisotopic values. +*) + +open BioFSharp +open BioFSharp.Mz + +let peptide = BioList.ofAminoAcidString "ANLGMEVMHER" + +let mono : IBioItem -> float = BioItem.monoisoMass + +printfn "residues: %i" (List.length peptide) +printfn "monoisotopic mass of Ala: %f" (mono AminoAcids.Ala) + +(** +```text +residues: 11 +monoisotopic mass of Ala: 71.037114 +``` + +## Generating the b series + +`Fragmentation.Series.bOfBioList` walks the sequence from the N-terminus and +accumulates residue masses. Each element of the result is a +`PeakFamily`, the shape introduced on the +[peaks and peak arrays]({{root}}01_01_peaks_and_peak_arrays.html) page: a main +peak tagged with its ion series flag plus a list of dependent loss peaks. The +masses are neutral fragment masses, no proton has been added yet. A small +printer makes the ladder readable. +*) + +let printLadder (prefix: string) (families: PeakFamily list) = + families + |> List.iteri (fun i family -> + let losses = + family.DependentPeaks + |> List.map (fun d -> sprintf "%A at %.5f" d.Iontype d.Mass) + |> String.concat " " + printfn "%s%-2i %A %10.5f %s" prefix (i + 1) family.MainPeak.Iontype family.MainPeak.Mass losses) + +let bSeries = Fragmentation.Series.bOfBioList mono peptide + +printfn "b families: %i" bSeries.Length +printLadder "b" bSeries + +(** +```text +b families: 11 +b1 B 71.03711 +b2 B 185.08004 B, lossNH3 at 168.05349 +b3 B 298.16411 B, lossNH3 at 281.13756 +b4 B 355.18557 B, lossNH3 at 338.15902 +b5 B 486.22605 B, lossNH3 at 469.19950 +b6 B 615.26865 B, lossNH3 at 598.24210 B, lossH2O at 597.25808 +b7 B 714.33706 B, lossNH3 at 697.31051 B, lossH2O at 696.32650 +b8 B 845.37755 B, lossNH3 at 828.35100 B, lossH2O at 827.36698 +b9 B 982.43646 B, lossNH3 at 965.40991 B, lossH2O at 964.42589 +b10 B 1111.47905 B, lossNH3 at 1094.45250 B, lossH2O at 1093.46849 +b11 B 1267.58016 B, lossNH3 at 1250.55361 B, lossH2O at 1249.56960 +``` + +The numbers check out by hand. b2 covers alanine and asparagine, and their +residue masses sum to 71.03711 + 114.04293 = 185.08004, exactly the printed +neutral mass. Protonation adds 1.00728, giving the singly charged ion at m/z +186.08732 that the peaks page constructed manually. + +The generator emits one family per residue, so the 11 residue peptide yields 11 b +families, and the last one spans the full sequence. The loss dependents +are cumulative: once the ladder has passed a loss prone residue, every longer +fragment carries that loss peak. The asparagine at position 2 puts an ammonia +loss on b2 and on every later b ion, and the glutamate at position 6 adds a +water loss from b6 on. b1 has no dependents because alanine triggers neither +rule. + +## Generating the y series + +`Fragmentation.Series.yOfBioList` builds the C-terminal ladder the same way, +except that the accumulator starts with the mass of one water molecule, because +a y fragment keeps the C-terminal hydroxyl and picks up a hydrogen at the break. +The function returns the ladder longest first, the head of the list is the full +length y11. For display we reverse it so that y1 comes first, and the code shows +that reordering explicitly. +*) + +let ySeries = Fragmentation.Series.yOfBioList mono peptide + +printfn "y families: %i" ySeries.Length +printfn "mass of the first returned family: %.5f" ySeries.Head.MainPeak.Mass + +let yAscending = ySeries |> List.rev + +printLadder "y" yAscending + +(** +```text +y families: 11 +mass of the first returned family: 1285.59073 +y1 Y 174.11168 Y, lossNH3 at 157.08513 +y2 Y 303.15427 Y, lossNH3 at 286.12772 Y, lossH2O at 285.14370 +y3 Y 440.21318 Y, lossNH3 at 423.18663 Y, lossH2O at 422.20262 +y4 Y 571.25367 Y, lossNH3 at 554.22712 Y, lossH2O at 553.24310 +y5 Y 670.32208 Y, lossNH3 at 653.29553 Y, lossH2O at 652.31151 +y6 Y 799.36467 Y, lossNH3 at 782.33812 Y, lossH2O at 781.35411 +y7 Y 930.40516 Y, lossNH3 at 913.37861 Y, lossH2O at 912.39459 +y8 Y 987.42662 Y, lossNH3 at 970.40007 Y, lossH2O at 969.41606 +y9 Y 1100.51069 Y, lossNH3 at 1083.48414 Y, lossH2O at 1082.50012 +y10 Y 1214.55361 Y, lossNH3 at 1197.52706 Y, lossH2O at 1196.54305 +y11 Y 1285.59073 Y, lossNH3 at 1268.56418 Y, lossH2O at 1267.58016 +``` + +The cumulative loss rule reads differently from this side because the ladder +grows from the C-terminus. The C-terminal arginine puts an ammonia loss on y1 +and with it on every y ion. The glutamate one position further in adds a water +loss from y2 on. y11 is the intact peptide plus water, and its 1285.59073 Da +match the 1285.59 Da neutral precursor mass determined on the charge state +page. + +The module offers the remaining series of the nomenclature through the same +pattern, `aOfBioList`, `cOfBioList`, `xOfBioList` and `zOfBioList`, plus +combined variants such as `abOfBioList` or `yzOfBioList` that return several +series in one list. + +## Matching the predictions against the measured spectrum + +The point of all this is that predicted fragments of the correct peptide should +be findable in the measured spectrum. We load the MS2 scan and look up every +predicted b and y ion in its m/z array as a singly charged species. +`Mass.toMZ` from BioFSharp does the charge conversion, it adds one proton mass +of 1.00728 per charge and divides by the charge. A predicted ion counts as +matched when a measured peak lies within 0.05 Da of it, and we keep the closest +such peak. +*) + +open BioFSharp.FileFormats.MGF +open BioFSharp.IO + +let ms2 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms2Example.mgf") + |> List.head + +let measuredMz = ms2.Mass + +let predictions = + (bSeries |> List.mapi (fun i f -> sprintf "b%i" (i + 1), f.MainPeak.Mass)) + @ (yAscending |> List.mapi (fun i f -> sprintf "y%i" (i + 1), f.MainPeak.Mass)) + +let matched = + predictions + |> List.choose (fun (label, neutralMass) -> + let predictedMz = Mass.toMZ neutralMass 1. + let close = measuredMz |> Array.filter (fun mz -> abs (mz - predictedMz) <= 0.05) + if Array.isEmpty close then None + else Some (label, predictedMz, close |> Array.minBy (fun mz -> abs (mz - predictedMz)))) + +printfn "matched %i of %i predicted singly charged ions" matched.Length predictions.Length +printfn "ion predicted measured" +matched +|> List.iter (fun (label, predicted, measured) -> + printfn "%-4s %9.4f %9.4f" label predicted measured) + +(** +```text +matched 18 of 22 predicted singly charged ions +ion predicted measured +b2 186.0873 186.0879 +b3 299.1714 299.1703 +b4 356.1928 356.1917 +b5 487.2333 487.2292 +b6 616.2759 616.2792 +b7 715.3443 715.3339 +b9 983.4437 983.4372 +b10 1112.4863 1112.5331 +y1 175.1190 175.1183 +y2 304.1615 304.1598 +y3 441.2205 441.2191 +y4 572.2609 572.2601 +y5 671.3294 671.3284 +y6 800.3719 800.3729 +y7 931.4124 931.4112 +y8 988.4339 988.4343 +y9 1101.5180 1101.5180 +y10 1215.5609 1215.5282 +``` + +18 of the 22 predictions find a measured peak, covering the y ladder from y1 to +y10 and most of the b ladder. The misses are plausible as well. b1 at m/z 72.04 +lies below the first recorded peak of the scan at m/z 100.67, and the other +three predictions simply have no peak within the tolerance. For a wrong +candidate sequence only a few predictions would hit a peak by chance. Turning +this observation into a score is the job of the scoring pages. + +## Generating a target and decoy pair + +Search engines need to know what a random match looks like, otherwise there is +no way to tell a convincing score from a lucky one. The standard trick is to +score every spectrum against the real candidate (the target) and additionally +against a deliberately wrong candidate of the same length and composition (the +decoy), obtained by reversing the sequence. The decoy scores calibrate the +score distribution of random matches, which later drives +[false discovery rate control]({{root}}04_01_fdr_control.html). + +`Fragmentation.Series.fragmentMasses` produces both at once. It takes the +N-terminal series function, the C-terminal series function, the mass function +and the peptide, and returns a `FragmentMasses` record with `TargetMasses` for +the given sequence and `DecoyMasses` computed from the reversed sequence. +*) + +let fragments = + Fragmentation.Series.fragmentMasses + Fragmentation.Series.bOfBioList + Fragmentation.Series.yOfBioList + mono + peptide + +printfn "target families: %i" fragments.TargetMasses.Length +printfn "decoy families: %i" fragments.DecoyMasses.Length + +fragments.DecoyMasses +|> List.truncate 3 +|> List.iter (fun f -> printfn "decoy %A ion at %.5f" f.MainPeak.Iontype f.MainPeak.Mass) + +(** +```text +target families: 22 +decoy families: 22 +decoy B ion at 156.10111 +decoy B ion at 285.14370 +decoy B ion at 422.20262 +``` + +Both lists concatenate the b and the y families, 22 entries each for our +peptide. The first decoy b ion weighs 156.10111 Da, the residue mass of +arginine, because the reversed sequence REHMVEMGLNA starts with the arginine +that ANLGMEVMHER ends with. + +## Converting neutral masses to charged m/z ladders + +The families so far carry neutral masses, while a spectrum shows charged ions. +In a real MS2 scan a fragment can appear at more than one charge when the +precursor charge allows it, our 2+ precursor can hand both protons to one +fragment, so the doubly charged ladder is worth predicting as well. +`Fragmentation.ladderElement` takes a peak family list and a charge list and +returns one family per input family and charge. Each resulting +`LadderedTaggedMass` carries the ion flag, the `MassOverCharge` value, the +`Number` of the ion within its series and the `Charge`. Internally the function +groups the input by ion type and sorts each group by mass, which is how the +number is assigned, so the mixed b and y list can go in as is. +*) + +let laddered = Fragmentation.ladderElement (bSeries @ ySeries) [1.; 2.] + +printfn "laddered families: %i" laddered.Length + +laddered +|> List.filter (fun f -> f.MainPeak.Number <= 2) +|> List.iter (fun f -> + let m = f.MainPeak + printfn "%A ion %i at charge %.0f: m/z %9.4f" m.Iontype m.Number m.Charge m.MassOverCharge) + +(** +```text +laddered families: 44 +B ion 1 at charge 1: m/z 72.0444 +B ion 1 at charge 2: m/z 36.5258 +B ion 2 at charge 1: m/z 186.0873 +B ion 2 at charge 2: m/z 93.5473 +Y ion 1 at charge 1: m/z 175.1190 +Y ion 1 at charge 2: m/z 88.0631 +Y ion 2 at charge 1: m/z 304.1615 +Y ion 2 at charge 2: m/z 152.5844 +``` + +The 22 input families become 44 laddered ones, one per charge. The 1+ values repeat +what the matching section used, b2 at m/z 186.0873 for example, and dependent loss +peaks are laddered alongside their main peak with the same number and charge. + +## Where the fragments go next + +The `TheoreticalSpectra` module pairs each candidate peptide coming out of a database +search with its predicted target and decoy spectrum, and a scorer such as +[SEQUEST-like scoring]({{root}}02_03_sequest_like_scoring.html) then quantifies how +well the measured peaks agree with each prediction. +*) From c212804c587bc092143b8ec2352143059b34f623 Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 21:50:12 +0200 Subject: [PATCH 06/11] Add peptide search database documentation --- docs/02_02_search_databases.fsx | 367 ++++++++++++++++++++ docs/data/Chlamy_Cp.fastA | 585 ++++++++++++++++++++++++++++++++ 2 files changed, 952 insertions(+) create mode 100644 docs/02_02_search_databases.fsx create mode 100644 docs/data/Chlamy_Cp.fastA diff --git a/docs/02_02_search_databases.fsx b/docs/02_02_search_databases.fsx new file mode 100644 index 0000000..de9c3a8 --- /dev/null +++ b/docs/02_02_search_databases.fsx @@ -0,0 +1,367 @@ +(** +--- +title: Peptide search databases +category: Peptide identification +categoryindex: 2 +index: 2 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "nuget: System.Data.SQLite.Core, 1.0.119" +#r "nuget: Newtonsoft.Json, 13.0.4" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(*** hide ***) +let dbFolder = System.IO.Path.Combine(System.IO.Path.GetTempPath(), "BioFSharp.Mz_docs_searchdb") +System.IO.Directory.CreateDirectory dbFolder |> ignore + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Peptide search databases + +Bottom-up proteomics identifies proteins through their peptides. The mass of an +intact protein is not specific enough to identify it, so the sample is digested +with a protease, almost always trypsin, and the mass spectrometer measures the +resulting peptides. Trypsin cuts C-terminal to lysine and arginine, except when +a proline follows, which yields peptides of around 14 residues on average that +ionize well. When the instrument selects a peptide ion for fragmentation, it +records the precursor m/z, and +[charge state determination]({{root}}01_03_charge_state_determination.html) +turns that into a neutral mass. Identification then becomes a lookup problem: +which peptides from the organism's proteome could have this mass? + +A run holds tens of thousands of spectra, and each one needs its candidate +peptides with every modification variant and a precomputed mass. The `SearchDB` +module does this preparation once. It digests the FASTA proteome, generates the +modified variants of each peptide, precomputes their masses and stores +everything in a SQLite database file with an index on the mass column. A search +engine then asks for all candidates within a narrow mass window around a +measured precursor and gets them back from an indexed query. + +This page builds such a database for the chloroplast proteome of +*Chlamydomonas reinhardtii* and queries it by mass. The same organism +accompanies the rest of these pages. + +## Describing the search space + +Everything the database will contain is decided up front by a `SearchDbParams` +record. The parameters are the identity of the database: two runs with the same +parameters mean the same database, and any change means a different one. + +A fixed modification is applied to every occurrence of its target +residue, the standard example being carbamidomethylation of cysteine from the +sample preparation. A variable modification may or may not be present, so the +database stores each peptide with and without it. The most common variable +modification is oxidation of methionine, which adds one oxygen atom. +`createSearchModification` describes it: a name, a +[Unimod](https://www.unimod.org) accession, a description, a flag whether the +modification is biological in origin, the composition as an elemental formula, +the target sites, whether the composition is added or subtracted, and a short +code that will mark the modification inside stored sequence strings. +*) + +open System.IO +open BioFSharp +open BioFSharp.Mz + +let oxidationM = + SearchDB.createSearchModification + "Oxidation'Met'" "35" "Oxidation of methionine" true "O" + [ SearchDB.Specific(AminoAcids.Met, ModificationInfo.ModLocation.Residual) ] + SearchDB.SearchModType.Plus "ox" + +let oxidationDelta = + SearchDB.massFBy SearchDB.MassMode.Monoisotopic (SearchDB.getModBy oxidationM) + +printfn "mass shift of %s: %.6f Da" oxidationM.Name oxidationDelta + +(** +```text +mass shift of Oxidation'Met': 15.994915 Da +``` + +The composition "O" translates to the monoisotopic mass of one oxygen atom, the +expected +15.995 Da shift. With the modification in hand we can assemble the +full parameter record. `createSearchDbParams` takes every field as a positional +argument. +*) + +let fastaPath = __SOURCE_DIRECTORY__ + "/data/Chlamy_Cp.fastA" + +let fastaHeaderToName (header: string) = header.Split('|').[1].Trim() + +let searchDbParams = + SearchDB.createSearchDbParams + "Chlamy_Cp_trypsin" // database name + dbFolder // folder for the db file + fastaPath // proteome FASTA + fastaHeaderToName // header -> accession + (Digestion.Table.getProteaseBy "Trypsin") // protease + 0 2 // min/max missed cleavages + 15000. // MaxMass + 4 40 // MinPepLength/MaxPepLength + [] // isotopic mods + SearchDB.MassMode.Monoisotopic // mass mode + (SearchDB.massFBy SearchDB.MassMode.Monoisotopic) // matching mass function + [] // fixed mods + [ oxidationM ] // variable mods + 2 // variable mod threshold + +printfn "database file name: %s" (Path.GetFileName(SearchDB.Db.getNameOf searchDbParams)) +printfn "protease: %s" searchDbParams.Protease.Name + +(** +```text +database file name: Chlamy_Cp_trypsin.db +protease: Trypsin +``` + +The name and folder determine where the SQLite file lands, here a folder under the +system temp directory, and `SearchDB.Db.getNameOf` shows the resulting file name. +`fastaHeaderToName` extracts a protein accession from each FASTA header line: our +headers look like `sp|P19528| cytochrome b6/f complex subunit 4`, so splitting at `|` +and taking the second field yields the accession. The protease comes from BioFSharp's +`Digestion.Table`, either by name as shown or directly as `Digestion.Table.Trypsin`. + +A missed cleavage is a lysine or arginine the protease failed to cut, and real +digests always contain such peptides, so search databases typically include peptides +with up to two or three missed cleavages, here up to two. + +One implementation detail matters when choosing `MinPepLength` and `MaxPepLength`: +the digest filter compares +`CleavageEnd - CleavageStart`, which is the peptide length minus one, strictly +against both bounds. The shortest stored peptide therefore has +`MinPepLength + 2` residues and the longest has `MaxPepLength` residues. With +the values 4 and 40 above, the database contains peptides of 6 to 40 residues. +`MaxMass` is recorded with the database and participates in its identity. + +Metabolic labeling such as full 15N would go into the isotopic modification list, +empty here, and the database would then store every peptide in a light and a heavy +form. `MassMode.Monoisotopic` together with `massFBy` selects the memoized +monoisotopic mass function used for all mass computations. Fixed modifications stay +empty. Oxidation of methionine goes in as a variable modification, and the threshold +of 2 caps how many variable modifications a single peptide may carry. + +## Building and connecting to the database + +`connectOrCreateDB` checks whether a database +file matching the parameters already exists. If it does, it simply reconnects. +If not, it reads the FASTA, digests every protein, generates the modified +variants, computes their masses and bulk inserts everything. The returned +value is an open `SQLiteConnection`. A few SQL counts show what the build +produced. +*) + +open System.Data.SQLite + +let cn = SearchDB.connectOrCreateDB searchDbParams + +let countRows table = + use cmd = new SQLiteCommand(sprintf "SELECT COUNT(*) FROM %s" table, cn) + cmd.ExecuteScalar() :?> int64 + +printfn "database file exists: %b" (File.Exists(SearchDB.Db.getNameOf searchDbParams)) +printfn "proteins: %i" (countRows "Protein") +printfn "distinct peptide sequences: %i" (countRows "PepSequence") +printfn "mass entries (ModSequence): %i" (countRows "ModSequence") + +(** +```text +database file exists: true +proteins: 74 +distinct peptide sequences: 6415 +mass entries (ModSequence): 9355 +``` + +The 74 chloroplast proteins digest into several thousand distinct peptides, +and the variable methionine oxidation expands them into more rows in the +`ModSequence` table, which holds one row per modified variant with its +precomputed mass. Calling `connectOrCreateDB` a second time with the same +parameters finds the parameter record stored inside the file and reconnects +without rebuilding anything. Changing any parameter, even only `MaxMass`, +changes the identity and triggers a fresh build. + +## Looking up candidates by precursor mass + +`getThreadSafePeptideLookUpFromFileBy` prepares the mass window query against +the open connection. The result is a function taking a lower and an upper +neutral mass and returning every `ModSequence` row in between. + +To query something realistic we first need a mass a spectrometer could have +measured. We digest the first protein of the FASTA in memory with the same +trypsin instance and pick the first tryptic peptide that fits the stored +length range and contains a methionine. Its neutral mass is the residue masses +summed up plus one water for the termini. +*) + +open BioFSharp.IO + +let firstProtein = + Fasta.read BioArray.ofAminoAcidString fastaPath + |> Seq.head + +let targetPeptide = + Digestion.BioArray.digest Digestion.Table.Trypsin 0 (firstProtein.Sequence |> Array.ofSeq) + |> Array.find (fun p -> + p.PepSequence.Length >= 6 && p.PepSequence.Length <= 40 + && List.contains AminoAcids.Met p.PepSequence) + +let targetMass = + targetPeptide.PepSequence + |> List.sumBy BioItem.monoisoMass + |> (+) (BioItem.monoisoMass ModificationInfo.Table.H2O) + +printfn "protein: %s" (fastaHeaderToName firstProtein.Header) +printfn "peptide: %s" (BioList.toString targetPeptide.PepSequence) +printfn "neutral monoisotopic mass: %.5f Da" targetMass + +(** +```text +protein: P19528 +peptide: LLGVLLMAAVPAGLITVPFIESINK +neutral monoisotopic mass: 2578.51720 Da +``` + +A production pipeline queries the database with a 30 ppm window around the +measured precursor mass. +*) + +let lookUpByMass = SearchDB.getThreadSafePeptideLookUpFromFileBy cn searchDbParams + +let ppmToDalton ppm mass = mass * ppm / 1000000. + +let showHits (hits: SearchDB.LookUpResult list) = + hits + |> List.sortBy (fun h -> h.Mass) + |> List.iter (fun h -> + printfn "%-30s mass %11.5f rounded %11i pepSeqID %i modSeqID %i globalMod %i" + h.StringSequence h.Mass h.RoundedMass h.PepSequenceID h.ModSequenceID h.GlobalMod) + +let tolerance = ppmToDalton 30. targetMass + +printfn "30 ppm at this mass: %.5f Da" tolerance + +let hits = lookUpByMass (targetMass - tolerance) (targetMass + tolerance) + +showHits hits + +(** +```text +30 ppm at this mass: 0.07736 Da +LLGVLLMAAVPAGLITVPFIESINK mass 2578.51720 rounded 2578517204 pepSeqID 4 modSeqID 3 globalMod 0 +``` + +The window returns exactly the peptide we computed the mass for, as a +`LookUpResult`. `StringSequence` is the stored sequence, `Mass` the +precomputed neutral mass and `RoundedMass` that mass multiplied by one million +and stored as an integer, which is the indexed column the BETWEEN query runs +against. `PepSequenceID` identifies the plain peptide sequence and +`ModSequenceID` the specific modified variant. `GlobalMod` tells whether the +entry belongs to the isotopically labeled form of the database. We configured +no isotopic modification, so it is 0 for every entry. The `BioSequence` field, +not printed here, holds the sequence parsed back into a BioFSharp amino acid +list, ready for fragment prediction. + +The oxidized form of the same peptide weighs one oxygen more and sits in its +own mass window. Querying 30 ppm around that mass returns the variant carrying +the modification. +*) + +let oxidizedMass = targetMass + oxidationDelta + +let oxHits = lookUpByMass (oxidizedMass - tolerance) (oxidizedMass + tolerance) + +showHits oxHits + +(** +```text +LLGVLL[ox]MAAVPAGLITVPFIESINK mass 2594.51212 rounded 2594512119 pepSeqID 4 modSeqID 4 globalMod 0 +``` + +The modification appears inside the sequence string as the `[ox]` code +directly in front of the modified residue, the same code we passed to +`createSearchModification`. Both variants share the `PepSequenceID` of the +plain sequence but have distinct `ModSequenceID`s and masses. A search engine +scoring this window would now treat the oxidized sequence as its own +candidate. + +## Mapping peptides back to proteins + +After spectra have been matched, the pipeline needs to know which proteins a +peptide belongs to, because the final goal is a protein list. The +`CleavageIndex` table stores this mapping, and +`getProteinPeptideLookUpFromFileBy` prepares a lookup from a `PepSequenceID` +to the accessions of all proteins containing that peptide, each accession +paired with the peptide sequence the ID stands for. It expects an +in-memory copy of the database, which `copyDBIntoMemory` produces from the +open file connection, so the many lookups of a full run avoid disk access. +*) + +let memoryDB = SearchDB.copyDBIntoMemory cn + +let proteinsOfPeptide = SearchDB.getProteinPeptideLookUpFromFileBy memoryDB + +let firstHit = hits |> List.minBy (fun h -> h.Mass) + +proteinsOfPeptide firstHit.PepSequenceID +|> List.iter (fun (accession, peptideSequence) -> + printfn "peptide %s occurs in protein %s" peptideSequence accession) + +(** +```text +peptide LLGVLLMAAVPAGLITVPFIESINK occurs in protein P19528 +``` + +The peptide maps back to P19528, the cytochrome b6/f subunit the FASTA starts +with, and the returned accession is exactly what our `fastaHeaderToName` +function extracted during the build. This reverse mapping is the raw material +for [protein inference]({{root}}04_02_protein_inference.html), where shared +peptides make the mapping ambiguous and need to be resolved. + +## Caching repeated lookups + +Consecutive precursors in a run often have similar masses, so their candidate +windows overlap and the same peptides would be fetched and their fragments +predicted again. The `Cache` module addresses this with a thin wrapper around +`SortedList`, keyed by the same rounded integer masses the database uses. +`getPeptideLookUpWithMemBy` combines such a cache with the database lookup and +fragment prediction, so a peptide that already went through the pipeline is +served from memory. Creating a cache and storing a result under its rounded +mass looks like this. +*) + +let lookUpCache = Cache.createCache list> + +Cache.addItem lookUpCache (firstHit.RoundedMass, hits) + +printfn "cached entries: %i" lookUpCache.Count +printfn "contains key %i: %b" firstHit.RoundedMass (fst (Cache.getItemBy lookUpCache firstHit.RoundedMass)) + +(** +```text +cached entries: 1 +contains key 2578517204: true +``` + +`addItem` inserts or replaces the value under a key and `getItemBy` retrieves it. +The memoized lookup itself is wired up by the search engine. + +The next step is to predict each candidate's fragments and compare them to the +measured spectrum, the subject of +[SEQUEST-like scoring]({{root}}02_03_sequest_like_scoring.html). +*) diff --git a/docs/data/Chlamy_Cp.fastA b/docs/data/Chlamy_Cp.fastA new file mode 100644 index 0000000..6cdd577 --- /dev/null +++ b/docs/data/Chlamy_Cp.fastA @@ -0,0 +1,585 @@ +>sp|P19528| cytochrome b6/f complex subunit 4 GN=petD PE=petD.p01 +MSVTKKPDLSDPVLKAKLAKGMGHNTYGEPAWPNDLLYMFPVVILGTFACVIGLSVLDPA +AMGEPANPFATPLEILPEWYFYPVFQILRVVPNKLLGVLLMAAVPAGLITVPFIESINKF +QNPYRRPIATILFLLGTLVAVWLGIGSTFPIDISLTLGLF* +>sp|P19529| photochlorophyllide reductase subunit B GN=chlB PE=chlB.p01 +MKLAYWMYAGPAHIGVLRVSSSFKNVHAIMHAPLGDDYFNVMRSMLERERDFTPVTASIV +DRHVLARGSQEKVVENITRKNKEETPDLILLTPTCTSSILQEDLHNFVESALAKPVQIDE +HADHKVTQQSALSSVSPLLPLEENTLIVSELDKKLSPSSKLHINMPNICIPEGEGEGEQT +KNSIFVKSATLTNLSEEELLNQEHHTKTRNHSDVILADVNHYRVNELQAADRTLEQIVRY +YISQAQKQNCLNITKTAKPSVNIIGIFTLGFHNQHDCRELKRLFNDLGIQINEIIPEGGN +VHNLKKLPQAWFNFVPYREIGLMTAMYLKSEFNMPYVAITPMGLIDTAACIRSICKIITT +QLLNQTATVQEPSKFIYPKATSLEQTNILETSQKETILKDNPDSGNTLSTTVEEIETLFN +KYIDQQTRFVSQAAWFSRSIDCQNLTGKKAVVFGDATHSAAMTKLLAREMGIKVSCAGTY +CKHDADWFREQVSGFCDQVLITDDHTQVGDMIAQLEPAAIFGTQMERHVGKRLDIPCGVI +SAPVHIQNFPLGYRPFLGYEGTNQIADLVYNSFNLGMEDHLLQIFGGHDSENNSSIATHL +NTNNAINLAPGYLPEGEGSSRTSNVVSTISSEKKAIVWSPEGLAELNKVPGFVRGKVKRN +TEKYALQKNCSMITVEVMYAAKEALSA* +>sp|P19530| photosystem II protein K GN=psbK PE=psbK.p01 +MTTLALVLAKLPEAYAPFAPIVDVLPVIPVFFILLAFVWQAAVSFR* +>sp|P19531| elongation factor Tu GN=tufA PE=tufA.p01 +MSRAKFERKKPHVNIGTIGHVDHGKTTLTAAITMTLAAAGGSVGKKYDEIDSAPEEKARG +ITINTAHVEYETEKRHYAHVDCPGHADYVKNMITGAAQMDGAILVVSGADGPMPQTKEHI +LLAKQVGVPNVVVFLNKEDQVDDKELLELVELEVRETLDKYEFPGDEIPVVPGSALLALE +ALIENPKTQRGENKWVDKIYQLMDNVDSYIPTPQRETDKPFLLAVEDVLSITGRGTVATG +RVERGALRISDNVEIVGLRPTQTAVVTGLEMFKKTLDETLAGDNVGVLLRGVQKKDIERG +MVIAKPGTITPHTKFEAQVYVLTKEEGGRHSAFMIGYQPQFYVRTTDVTGKVVGFNHIQM +RNPSSVAEEHSNKMAMPGDRISMTVELINPIAIEKGMRFAIREGGRTVGAGVVTNIVQ* +>sp|P19532| ribosomal protein L20 GN=rpl20 PE=rpl20.p01 +MTRVKRGNVSRKRHKKILNMSKGFRGAASTLFRTANQQNMKALRYSYRNRRQKKRDFRRM +WITRVNSAVRRYGLNYSEFMNYLKTHKIQLNRKVIAQLSICDPEAFMQLLLF* +>sp|P19533| ATP-dependent Clp protease proteolytic subunit GN=clpP PE=clpP.p01 +MPIGVPRIIYCWGEELPAQWTDIYNFIFRRRMVFLMQYLDDELCNQICGLLINIHMEDRS +KELEKKEMEKSGLFKSGTAKTKGKDTVKKENLSGGASAKRQSVEDLLTSDNDFGIEENHL +LEQYTLQKITTEWLNWNAQFFDYSDEPYLYYLADILSKDFSPNQDKDSANLNFAKSSANK +QAFQNPAEMTKLIKNLKNLKNFSTGSKNVKQNLDVYSPFRLLANFAPQNYNLEHPNQNLA +EIYSLLKTSTQNTNQPFTKKLIDNLSHKELMNRLQSPEKLVASSEKALGQRRLKQRYVQE +RLGSGGLSNSKALKAYNYLDQGALNNESGRSLYRKQTERVIQEEESKKVFMIINSFGGSV +GNGITVHDALQFIKAGSLTLALGVAASAASLALAGGTIGERYVTEGCHTMIHQPEGGLNG +QASDIWIDSQEIMKIRLDVAEIYSLSTYRPRHKILRDLDRDFYLTAMETIYYGLADEIAT +NEVMHSIVEMTNQVWSYHDSKQERLLESRASLVGDSTQTQESNS* +>sp|P19534| cytochrome b6 GN=petB PE=petB.p01 +MSKVYDWFEERLEIQAIADDITSKYVPPHVNIFYCIGGITFTCFLVQVATGFAMTFYYRP +TVAEAFASVQYIMTDVNFGWLIRSIHRWSASMMVLMMVLHVFRVYLTGGFKRPRELTWVT +GVIMAVCTVSFGVTGYSLPWDQVGYWAVKIVTGVPDAIPGVGGFIVELLRGGVGVGQATL +TRFYSLHTFVLPLLTAVFMLMHFLMIRKQGISGPL* +>sp|P19535| photochlorophyllide reductase subunit L GN=chlL PE=chlL.p01 +MKLAVYGKGGIGKSTTSCNISIALAKRGKKVLQIGCDPKHDSTFTLTGFLIPTIIDTLSS +KDYHYEDIWPEDVIYGGYGGVDCVEAGGPPAGAGCGGYVVGETVKLLKELNAFFEYDVIL +FDVLGDVVCGGFAAPLNYADYCIIVTDNGFDALFAANRIAASVREKARTHPLRLAGLIGN +RTSKRDLIDKYVEACPMPVLEVLPLIEEIRISRVKGKTLFEMSNKNNMTSAHMDGSKGDN +STVGVSETPSEDYICNFYLNIADQLLTEPEGVIPRELADKELFTLLSDFYLKI* +>sp|P19536| ribosomal protein L36 GN=rpl36 PE=rpl36.p01 +MKVRASVKKMCDNCRVIKRKGKVMVICSNAKHKQRQG* +>sp|P19537| ribosomal protein L23 GN=rpl23 PE=rpl23.p01 +MLDLVKYPVITQKTYIALFKDRQYTFDVDLRLTKPQIKKVFETLFNVDVISVNTHIPPRQ +KIRVGLAQGYRPRYKRAIITLKEGQSINYSLKNDN* +>sp|P19538| ribosomal protein L2 GN=rpl2 PE=rpl2.p01 +MGIRFLQAYTPGTRNRSVSDFSELTDKNSTPEKALTVSLHRAKGRNNRGIITCRHRGGGH +KRLYRQIDFRRDKIGVTAKVVRIEYDPNRNARIALLRYEDGEKRYIIHPRGLNIGDIIQS +DLNAPILIGNSLPLRNIPLGAEVHNVEFQPGSGGQLARSAGAMVEILAKEGNFVTIRLPS +KEIRLVSKNCWATVGQVGNIEAYNLTIGKAGRTRWLGKRPTVRGSVMNPVDHPHGGGEGR +APIGRSRPVTPWGRPALGQLTRKPKKYSNTLIVKKRKK* +>sp|P19539| ribosomal protein S19 GN=rps19 PE=rps19.p01 +MSRSLKKGPFVADHLLKKIEKLNAKGKKVVIKTWSRSSMIVPPMIGHTIGVYNGREHIPV +FVSDQMVGHRLGEFSPTRTYRGHAKKDKKAKR* +>sp|P19540| ribosomal protein L16 GN=rpl16 PE=rpl16.p01 +MLSPKRTKFRKPHRGHLRGKATRGNKIVFGDFALQAQEPCWITSRQIEAGRRVLTRYVRR +GGKLWIRIFPDKAVTMRPAGTRMGSGKGAPDYWVAVVHPGKILYEMQGVSETIARQAMRI +AAYKMPVKTKFLTKTV* +>sp|P19541| ribosomal protein L14 GN=rpl14 PE=rpl14.p01 +MIKPLSYLNVADNSGARELMCIRALGGSYRESANIGDVIIAVVKDALPNMPVKRSDIVRA +VIVRTRKGIRRENGMAIRFDDNAAVIINKEGNPRGTRVFGPIARELRDKNFTKIVSLAPE +VL* +>sp|P19542| ribosomal protein L5 GN=rpl5 PE=rpl5.p01 +MTQRLKNLYTKTIVPKLTTNFNYSNMHEVPKIEKIVINRGIGDASQNQKIVESSLKELAM +IAGQKGVVTRSKKAIAGFKLRQQMPVGVTVTLRGDRMYGFLDRLIHLALPRVRDFQGISS +KSFDKKGNYSLGLEEQLMFPEIEYDKIDQVRGMDISIVTTAKTQEEGLALLKEFGLPFK* +>sp|P19543| 30S ribosomal protein S8 GN=rps8 PE=rps8.p01 +MTITKTATLKQTMKSHGLINDSIGDMLTRIRNACLAKKSSVSIPFTRLNQNIAQILEQEG +FIQTYQVSLDSQDLTIRLKYRSKKIYRGKKKESCITNLKRISKPGLRIYSNHKDILRILG +GTGIVIVSTPEGLMTDREARLRGIGGELLCSVW* +>sp|P19544| photosystem I P700 chlorophyll a apoprotein A1 GN=psaA PE=psaA.p01 +MTISTPEREAKKVKIAVDRNPVETSFEKWAKPGHFSRTLSKGPNTTTWIWNLHADAHDFD +SHTSDLEEISRKVFSAHFGQLGIIFIWLSGMYFHGARFSNYEAWLSDPTHIKPSAQVVWP +IVGQEILNGDVGGGFQGIQITSGFFQLWRASGITSELQLYTTAIGGLVMAAAMFFAGWFH +YHKAAPKLEWFQNVESMLNHHLGGLLGLGSLAWAGHQIHVSLPVNKLLDAGVDPKEIPLP +HDLLLNRAIMADLYPSFAKGIAPFFTLNWSEYSDFLTFKGGLNPVTGGLWLSDTAHHHVA +IAVLFLVAGHMYRTNWGIGHSMKEILEAHRGPFTGEGHVGLYEILTTSWHAQLAINLALF +GSLSIIVAHHMYAMPPYPYLATDYGTQLSLFTHHTWIGGFCIVGAGAHAAIFMVRDYDPT +NNYNNLLDRVIRHRDAIISHLNWVCIFLGFHSFGLYIHNDTMSALGRPQDMFSDTAIQLQ +PVFAQWIQNTHFLAPQLTAPNALAATSLTWGGDLVAVGGKVAMMPISLGTSDFMVHHIHA +FTIHVTVLILLKGVLFARSSRLIPDKANLGFRFPCDGPGRGGTCQVSAWDHVFLGLFWMY +NSLSIVIFHFSWKMQSDVWGTVTASGVSHITGGNFAQSANTINGWLRDFLWAQSSQVIQS +YGSALSAYGLIFLGAHFVWAFSLMFLFSGRGYWQELIESIVWAHNKLKVAPAIQPRALSI +TQGRAVGVAHYLLGGIATTWSFFLARIISVG* +>sp|P19545| ribosomal protein S4 GN=rps4 PE=rps4.p01 +MSRYLGPRLRVIRRIGKLRGFTRKKPFRRVFKGFGGFKGKVIPPGQHGLTKLLKTRPYDS +SESDYLIRLKVKQRLRFNYGITERQLVNYVRKAKKIKESTGQVLLQFLEMRLDNIVFRLN +MAPTIPAARQLISHGHIRVNNKKVNIPSYMCKPKDVISVAMKQRSLQLVNKNLQEYYRRM +RFYKKRLEKTLPFILLKIKPLGLTSVTAAVELITKGNVRVNNKSVKTPNYICRPRDTVSL +RTKQGIKKVFLKNYLKG* +>sp|P19546| photosystem II protein D1 GN=psbA.1 PE=psbA.p01.1 +MTAILERRENSSLWARFCEWITSTENRLYIGWFGVIMIPCLLTATSVFIIAFIAAPPVDI +DGIREPVSGSLLYGNNIITGAVIPTSNAIGLHFYPIWEAASLDEWLYNGGPYQLIVCHFL +LGVYCYMGREWELSFRLGMRPWIAVAYSAPVAAASAVFLVYPIGQGSFSDGMPLGISGTF +NFMIVFQAEHNILMHPFHMLGVAGVFGGSLFSAMHGSLVTSSLIRETTENESANEGYRFG +QEEETYNIVAAHGYFGRLIFQYASFNNSRSLHFFLAAWPVIGIWFTALGLSTMAFNLNGF +NFNQSVVDSQGRVLNTWADIINRANLGMEVMHERNAHNFPLDLASTNSSSNN* +>sp|P19547| photosystem II protein D1 GN=psbA.2 PE=psbA.p01.2 +MTAILERRENSSLWARFCEWITSTENRLYIGWFGVIMIPCLLTATSVFIIAFIAAPPVDI +DGIREPVSGSLLYGNNIITGAVIPTSNAIGLHFYPIWEAASLDEWLYNGGPYQLIVCHFL +LGVYCYMGREWELSFRLGMRPWIAVAYSAPVAAASAVFLVYPIGQGSFSDGMPLGISGTF +NFMIVFQAEHNILMHPFHMLGVAGVFGGSLFSAMHGSLVTSSLIRETTENESANEGYRFG +QEEETYNIVAAHGYFGRLIFQYASFNNSRSLHFFLAAWPVIGIWFTALGLSTMAFNLNGF +NFNQSVVDSQGRVLNTWADIINRANLGMEVMHERNAHNFPLDLASTNSSSNN* +>sp|P19548| hypothetical protein GN=ycf12 PE=ycf12.p01 +MNIELALTLVSLVLVVSAGPLVVVLLSARGGNL* +>sp|P19549| ATP synthase CF1 epsilon subunit GN=atpE PE=atpE.p01 +MSLQISILTPERPFWNGQADEIILPTETGEMGVLKNHAPIITGLDVGAMLIRGGQASGSK +DEWNSYAIMGGFALVKQNQVTILANEAVSAENINPEEAKDAFETAKANLEKAEGVKEKVE +ANFAYKRAKARYQVVKVLKKI* +>sp|P19550| ribosomal protein S7 GN=rps7 PE=rps7.p01 +MPRRPINKKRTLLPDPVYNSVSVHMLVNRVLKSGKKSVAYRIVYNALKEIGDVTQKNPVE +VFEKALDNVTPRVEVKPRRRAGAIQMVPRVLRLGDRARANSLRWIMEACDKRSGQPMVTK +LKSEILDAYKKTGFAIRKKEELHKIAIANAMYAKKPQVIINAINLLVD* +>sp|P19551| ribosomal protein S14 GN=rps14 PE=rps14.p01 +MAKKSMIQRELKRQKLVMKYATKRAALKEQIKQTTFLKEKLSLHRKLQQLPRNSSAVRLH +NRCMITGRPKGYFRDFGLSRHVLREMAHQGLLPGVCKSSW* +>sp|P19552| photosystem II reaction center protein M GN=psbM PE=psbM.p01 +MEVNIYGLTATALFIIIPTSFLLILYVKTASTQD* +>sp|P19553| photosystem II protein Z GN=psbZ PE=psbZ.p01 +MTSILQVALLALIFVSFALVVGVPVVFATPNGWTDNKGAVFSGLSLWLLLVFVVGILNSF +VV* +>sp|P19554| cytochrome c biogenesis protein GN=ccsA PE=ccsA.p01 +MNFVNLEQIENSLRNATFCMLFLTTFLYWFYTAFYSTNPQQIINPLSLTNIKTNFTYPDG +FQANNINVSTTYLPINPVLNTEREEQPEANGTNGLLGVSSLVVNLKSIAIPRIMMGVSNL +LLVLLLLVRWEKSGHFPLSNLYESLMFLAWCCTFLYLLYCTSFTLLVEKMLGSLIAPCSL +LMNAFATFSLPKEMQQASPLVPALQSNWLMMHVTVMIISYATLIIGSLLSILFLILFKHK +KGTPKKYDNFINNLDALSYRIIGLGFPFLTIGILSGAVWANEAWGSYWSWDPKETWALLT +WLVFAIYLHTRLTKGWEGEKPAIIAAVGFLVVWFCYLGVNLIGEGLHSYGFFN* +>sp|P19555| photosystem II protein H GN=psbH PE=psbH.p01 +MATGTSKAKPSKVNSDFQEPGLVTPLGTLLRPLNSEAGKVLPGWGTTVLMAVFILLFAAF +LLIILEIYNSSLILDDVSMSWETLAKVS* +>sp|P19556| photosystem II reaction center protein N GN=psbN PE=psbN.p01 +MESPAFFFTFFLWFLLLSVTGYSVYVSFGPPSKKLRDPFEEHED* +>sp|P19557| photosystem II protein T GN=psbT PE=psbT.p01 +MEALVYTFLLVGTLGIIFFSIFFRDPPRMIK* +>sp|P19558| photosystem II 47 kDa protein GN=psbB PE=psbB.p01 +MGLPWYRVHTVVINDPGRLISVHLMHTALVSGWAGSMALFEISVFDPSDPVLNPMWRQGM +FVLPFMTRLGITQSWGGWTISGETATNPGIWSYEGVAAAHIILSGALFLASVWHWTYWDL +ELFRDPRTGKTALDLPKIFGIHLFLSGLLCFGFGAFHVTGVFGPGIWVSDPYGLTGSVQP +VAPSWGADGFDPYNPGGIASHHIAAGILGVLAGLFHLCVRPSIRLYFGLSMGSIETVLSS +SIAAVFWAAFVVAGTMWYGSAATPIELFGPTRYQWDQGFFQQEIQKRVQASLAEGASLSD +AWSRIPEKLAFYDYIGNNPAKGGLFRTGAMNSGDGIAVGWLGHASFKDQEGRELFVRRMP +TFFETFPVLLLDKDGIVRADVPFRKAESKYSIEQVGVSVTFYGGELDGLTFTDPATVKKY +ARKAQLGEIFEFDRSTLQSDGVFRSSPRGWFTFGHVCFALLFFFGHIWHGARTIFRDVFA +GIDDDINDQVEFGKYKKLGDTSSLREAF* +>sp|P19559| RNA polymerase alpha subunit GN=rpoA PE=rpoA.p01 +MTIYPNLKKIMTKTQSTDFFIACKESRIENNTNFYGCFYLGPFDESLSQTLANDLRRTLL +SELTGLAITSIEIEGVLHKFSTLTGMKEPVLDLICNLQNIVLRKETISSTNMNYRATKKT +YIGFLSVNGPRVIKAADLKLPAGLQCVDPNQYIATLAEDGFLNMKFNINEGKNYIKQKPY +NLDVTTLKKRNILLQNFKNKIGLTALKNKQLMSTTMEGKPLLSNSDNLRFKKSFKRMLTS +PNQSLKNKTSLGHDTVSNPIPLDAVFMPVTKINCIIEENNVYSDFSTDPSLEFSTHLVPS +LTTQTNLRQESTHKFVNKANSLLQENNLFRSEKVYLPNIYIPEGEGDALSLKGVSPYSDF +KTFLSTLNYNSLYQTSLFLNQSLGQNKLLPWQANTLFFDVTNFADSQSNDVNMNMDLAGD +KTSVQKIHSTGISNTDAQLNKLSLTKIKTFLGSSDKLQNKTYQSFLQPKYASSNLRTLLT +QRSLQKNQSVFMHSFLDDQAKHKELTSNTLRANKFQVMKTVVSIPPKTFKTNPINTFHNQ +SLTFEYAKNFKLKPLRKKSQLILEIWTNGSIHPRKALYQALIFLSNNFLKLQTVKMLGSM +FKSDLAYGNLKHSITNNYTYYQTNNLSQQSKKTLFKKAEAAASKQALNFLKPSILNTNNL +PGGTMDLYLQSSLNAPIGILKISLRSYTALKKAGITTLKDLIKYSKKDLFAIKNLGQKSV +FEIEQNLALLGLTLVTD* +>sp|P19560| ribosomal protein S2 GN=rps2 PE=rps2.p01 +MLNKKPPYLILSLIMIKTQKSKQTAVRKLIPGNVISLKITAMGANNVGINEFTFGIPVLV +PNAKLGETVQAKVLKILASKKIAIAKLIKVVTKTNATETNNLAALTPGATLDVTITKLGP +NSTGIADLGNNYNNVNKLIVKKSAITIGEKVTVLVTRVKNGYAFGVATVSTQRLNQVSTS +LTQNSFKGTKFTVVLPKNAKRYLKHLVFKVTTATAQNLSVNGFEQTLFKKGAVGLDTRIN +NQNGSAGNLTTVPTNTILFVKPNLGAKLGDKVQIQIIKFASIENGTLTYNVAIAKIIKLN +PLSTPQKKAFVRSSLRQMLKSGMHYGEKAIKCNARMKNYVWTRKKGTDTKVEARPLIKKG +RNLINLLKTRRCLTKALAQLTKYAAKGKTFLFVGTKKAASGLVARAALFSKKAFFVNTRW +LGGMLTNWKTILKSISKIRPILKEKQMIIKDILEKRQTIKARLIQKALLLRKKSKLMLKK +GRLLIQMLKQNNSNSTSGKQAVRFLFTEKTNLLNTKRKEFVSKGILLLEKRQQLVVKRQE +LITQSQTLKSKAIQLTNTYRNLLNNLICSRKKLRELKALLLVSHELYLFKQQAKQDNQNL +YMVSYNKFKTLNSDYILSNPPKEILNKMVSIIKGQGLVIKNNNLNLKTANNAKTLILSQL +LSKFSLFVPTIKTSINNLQNYISTQKTALNKVLALLNVVKTKMNVYVTLKTKLVAELRQI +KQTLQTERNIIRVLRRKLKQIAAQKRFIKFLPKLRYLPTPVTKIEQTARFLVKKFVDPKM +KYPMDSIYDKKLSRQSKKVAASRKKKWQRLEKYLGGISNMTKIKEKQIANNVAIIIGQQE +EMNAVRECQKLGIKMFHIVDTNCNPGLADHFIPANDDARNSIKFILGKFLTRIRLAHKIK +VKFKKTSLKK* +>sp|P19561| ribosomal protein S18 GN=rps18 PE=rps18.p01 +MRGLKSNKPKPRIQKVSVRKFRRKVLSLSQILSRLRQKRNQKIEQQKRNKPIKPLIPPKT +LVIVLKEKPEKTLYNRRYIDYKHSGLLQRYIGLGGKILPRRQTRLTAKQQRFVAKTIKTA +RVMGLLPFVSKERSFFK* +>sp|P19562| photosystem I assembly protein Ycf4 GN=ycf3 PE=ycf3.p01 +MTQNNILIRRYIIVGERRFSNYWWAIVIFLGSCGFLATGICSYLGIPNWLSLLNIGTTFS +SETETLASGIVPFFPQGLLMSFYGSLGFLLSIYWSLLIFWNVGGGFNEFNKKEGFVRIFR +WGYPGKNRKIDLSYSLKDIEAIRVELKQGLDAQRTIYLRLKGKREIPLTGIGQPLTLKEI +EKQASELANFLQVSLEA* +>sp|P19563| 30S ribosomal protein S9 GN=rps9 PE=rps9.p01 +MAILAKAVGRRKEAVAQVQIKEGNGLFIINNKSAQEYLHNDFYSLLAVKAPFDVLSTSKK +ADMVSPDLSLTNLESTFTNLMQFSGGENAVKFDTIVKVKGGGLMGQTEAIRLGISRALCL +LSTNTNPSANQNNLPNIYIPEGEGEALAIPNASDIRKQLKDKGYLTQDSRVKERRKYGLK +KARKASQYHKR* +>sp|P19564| photosystem II protein V GN=psbE PE=psbE.p01 +MAGKPVERPFSDILTSIRYWVIHSITVPALFIAGWLFVSTGLAYDVFGTPRPNEYFTEDR +QEAPLITDRFNALEQVKKLSGN* +>sp|P19565| RNA polymerase beta subunit II GN=rpoB-2 PE=rpoB-2.p01 +MTYSILTESIYNFKKKSSLDMQFLSPKNFQIENFTKIHDQTLPLKPFKTNRTAVAVVPPL +PIFSPELYKRQNKLKKTQNATIFSMGTSAFKNQVKYNLETYHRSNQDTCLIHKPAVKEGD +WVEVGDLLADSASSIGGELAIGHNIIVAYMPWEGYNYEDAILINERLVYEDIYTSIHIER +YEVTTKETKLGFEQITREIPDISENEIKHLDKTGIAKIGSWVEEGDILVGKITPFNIKTL +TPQQKLLYKIFDKQLSTTKDSSLRAPKGIKANVININILARQKIQINTKSKNTGKGSKPP +RASKAQNTMVSQPSYIHIYLAEKRKMQVGDKMAGRHGNKGIVSRILPRQDMPFLPDGAAV +DIVLNPLGVPSRMNVGQIYECLLGLAGRYLGEHYKIPPFDEMYGADASRSFVLSKLYEAR +KKTGLKWLLDPNHPGKIRLFDGRNSECFDQTVTVGIAYVLKLVHMVDDKMHARSTGPYSL +VTQQPLRGRSKQGGQRLGEMEVWAIEGYGAAFVLSEMLTIKSDDMTGRQNLWKNLIENKE +ISLGSPESFKVLICELQALCLDIGLFRKNKETSLPYFKMPGEESKTNVNANLTGVQSSKK +LEANSQTVKNSSMPNLVEIDNLLNLA* +>sp|P19566| RNA polymerase beta subunit I GN=rpoB-1 PE=rpoB-1.p01 +MLNISTNMMINQTLNFDSNQESQKNKTERISKTKLTRELITKTDHDIYVDDISFLQREDN +VNVSQIDLTARGVLNPEDQGVLHKSTEFGVFNKNKTGLPASVDLKIKQNSFFSVKQPDFN +KYLISDFVEIQRNSFFTLLEKGIIEEFSKRNPITNSKKTMEIFFYPDYYQLTPPEYSPSQ +AIIKSKSYTSKLYIPVQLTDKSKNIIKLKWVYIGDIPLMTKRGHFILNGCARVIVNQMVR +SPGIYYQKKIYENFSNKWSEKPENTFTRYFADLICNRGTWLRIEMDKYNKIWAQMKRVPK +IPIMWFLIAVGLTDKIVLKTVMDSKILLYNLDEDPLNPRKKPLPYVKTPPAAWSAIYNIV +FAKKIKAQEKAKKMLELTEGNPSSKSQTKNKTSASKKSKTLNVANTKGKTPAENIKTLSE +LIDLEKALFLKSEQGRKWIFNKFMNPRTYDLGKVGRVNFNRKLKLSISQDITTLTPQDLL +AATNNLILVSKGLRELDDIDHLKNRRVRTSGELIQIQIGVGLVRLEKTIREKMTYASGLS +SLPSQKFAFRSSKQRNQSPVGDAENATQLTIGNLINTKPFNGALREFFGTSPLSQFMDQI +NPLAELTHKRRLSSMGPGGVTRDSATLAIRGIHPSHYGRICPVETPEGKNTGLVNSLTAY +ARVNAAGYIETPFYRVYKGQVQKKTGLYFFSAKQEEKIKLGAPDLYTSEIGFLPKASIPV +RIVEDFTKISRNEIQYVGVAPIQMISIATSLIPFFEHDDANRALMGSNMQRQAVPILKPQ +RPIVGTGLEARAVSDSGHVITAKSSGIVMYTSSKEIIIYSLQ* +>sp|P19567| photosystem II protein VI GN=psbF PE=psbF.p01 +MTTKKSAEVLVYPIFTVRWLAIHGIAVPTIFFLGAITAMQFIQR* +>sp|P19568| photosystem II protein L GN=psbL PE=psbL.p01 +MARPNPNKQVVELNRTSLYWGLLLIFVLAVLFSSYIFN* +>sp|P19569| cytochrome b6/f complex subunit 5 GN=petG PE=petG.p01 +MVEPLLCGIVLGLVPVTIAGLFVTAYLQYRRGDLATY* +>sp|P19570| ribosomal protein S3 GN=rps3 PE=rps3.p01 +MGQKVHPLGFRVGITKKHQSQWFARFQKYAYSQSVFEDHMLRTTLVNLFSNLEKESALAT +KQSKNRGATQPKAPKITQIKIERGLIPYEIGIQIHSNDCLSITKAIDNIKVSKDLVTNLQ +KTRKYLFKAGTQLKNASMQVSDNLNVAEGENSTMLKTKTSASGTSVLKKRTFKIQTQKPT +KGKFVFKGKRKQKKKLSKAVFMRLKNIKRRFKKRQTIKKRYLNIISKGLLIRKKGNLIIR +NVKIKRKNKTARLTSRKSQPTLRSGSNLRDNLAPVKSKMQRFNNRMSKKFANLFLTKLNK +QFLVRLKAIMKFWHNQNVTKAPLGYNKKWSLAKSYALINNLKAKYLAKDILSLGSLRVQK +LRKLISILEKKSLVKMETLRKDFITFGTLSKTRAFGYYQMITFLKQLKELVTKIKKQTIA +NVTTKQESVVDNKLASNKTKIQNLIRAKSKQTKSITQKVVNNFVKLVDDNQAMANESRKI +KWISYLKDLVNKHRTENIFYYLATIATARKDLNALKRYTKQHANFLFGVNVENAKENPNA +LLQRVTKTLTQYSKNPLVNNDFENAEGLTKLQTAFLTQIESQRKMYKANLALTPKISIKF +FSVKTTNLLEKASTVADSIVDALEKRKAFRGVIKKAKEDLMLRSRVTRVKGVKIQVAGRL +NGAEIARSEWVRAGRVPLQTLRANIDYAYRTANTIYGIIGVKVWIFKGYSKI* +>sp|P19571| RNA polymerase beta' subunit GN=rpoC2 PE=rpoC2.p01 +MNIFFQRKLVKYFGNIKKKKHYLTVSEECNHYVFVKKTYVSSVFLKSLNKVNKTCNTNSG +ILYNQYSLGYINMHKQLPSLIRNKSKTIGNYGVAHPSRLVKCLVTKNATAPFFNFTFDKG +RLKNLVSWTLENYGQYKTVELLEQLKKTGFEYATKAGISLGLDDLKIPPKKKILLLEAEQ +LTKLTIHQYQRGDITAVERFQRLIDTWHRTSEQLKQEVINYFEETDILNPVYMMAFSGAR +GNISQVRQLVGMRGLMSDPQGQIIDFPIQSNFREGLTLTEYIISSYGARKGIVDTALRTA +NAGYLTRRLVDVAQHVIISHYDCGTHKGIFLTDMKEGNKTIVSAQSRIIGRVLARDIYKP +NSTITIAKRNQEISTDVAFEIGKVTNRIFVRSALTCNTTKLLCQLCYGWSLAQGNLVSVG +EAVGVIAAQSIGEPGTQLTMRTFHTGGVFSGDVSDEIRAPYNGFVYYDNKIPGILIRTLD +GKILFLTKSEGTLIFTADPNFNKPQALTDSFLNSGHGEKEKYEIKKYKIPAYTLLFIRNG +ESVLQKQVLAQITMISTKPNMRDTAELVIKAELEGLFYAKNLQVQKKILGPKPKFIGEGK +QNVLLDPKAMEIIVKARGWNFAWVLSGKRYEFPLLLKSFASSGDLITPNTIMAKHNLQLS +SPFLNVGTALQVGNSTDIPKLPLLGATSKFRAMSGVLTSSIVRRYPTKINSSSRPKVNQF +GKSNLFYTQVLQNKVNALNGEVDSYSETAKLPYWKNLNLKFVQKLNLSKFFKQYNLKNKT +KTKLAFYSLVQAKPLQASKLDSFKANTFQSKQLGRPIFIYRHCLLTKESNKKTVKLIHNI +QLQQSVLFLKIKKIKFYKMGYFQLVNSNKTAFLVSLSHNKNRLTLYHNKFNYNNSYNDII +VLPTSLTQDISSKKQIALKRFKPAYNLFQWFYPSLIKPSSKEAQNLTVQQVEFFNAREQM +HFNSRYSKTDFVQLLQNPFKLDFNKALHRPLCISESYKEIPTGEVSISPQNSVNLPMNTF +FNTDGVPDKKPNYTNMYAFWLKQQVLKSYKKRYKKYKLTSILKSHLSDKIYLPASVELQH +QSQQNNLLSLTKEFKALNSSKYLKNNQLQSRPLLYKQEKLLKTLVLKKWFKSNLLYVNSA +ISKEENMDGSSTQMSAHSLPKRQRKGAKVSKINRFLKIDNYIKQLTYFKTSKIMRIKHSF +NSLRVDFKFTSSKLVCPKRKRNLQIVCTLRGEEQTSVRESSVTHVNNLICSSKYKRSIRL +QNDSHLFTTTKKAKVPFTGYSDSHQAGKVLQSSQISSTKPKTQSSFTFFEYFKLKLAQSG +TKAILKHFKSLKAVNISKVSMDQSRNKGFNNWGKSSSECLNTSLGDKQALYFKTKTEANL +QLLTLVLENFYRKKQFANLLAKNLTILNKNLKLKKNHVHFLLYYYNKLSVQKPNAIFLLT +HMLAKANKIQISYSYVVPTSPKSKLNLDRGNLTTTNQKNVASLGGYQPASAGTKYYSEGI +RNRTRKNNKSSMTKITKIFNQLKLDTQFVIILLSPHKLLHHSHQLPQNTLGIYDKFNSNF +EMAKVTLLTHPLWFNQFQMKSIKEVGNKFINTRNNSLLENYVILLLSNNYVFNFLSIKSE +QINLPEEQESNLSLQEYKSPANCQEKALPSDNKKIIFDLKKLQHIVLTEQYKNNLRWVKN +KKPRFLPKDIDINTLEVNLDRIKPKNKLSNLNPTQQLQLSNSLLEFVKTIKINPTDLAKV +YSGSANTVMSIENCQPSLEVSNTFFLKTQKLLKLINKTKHSIALNQTKFINTSLLKGHLV +AYARPVFIITNKAEPFIAKQKKDLLLLPKNATINVLIKPQQEKNSLNKAIFPLGGNAANN +HSIFVSPNTNKLANPNVSYLKKHIYFNQMPFFDSPFRNASYLFSYTENSIKPLTKVDFMH +SFAQKNHKLLPESEREKRLQFKALHIPKQPCLNICFKSNSNLIFNSKTTNSLVIRKTTTN +YKYNIDLSEGVDFKKLKTNMFSARKCNNFKLDTALESKLFKGRPTLLNYKNVVTQTNYFS +PFEGELLATKTYKNYLDLNPYQSLKVGLMQLNTSSALATLVPKTKRAGNKSSKQKIKLNQ +GELSTELGSTIPQSGKHKTKTEKMRMAMFKYYLKTINSQKIIGNKGWSRFNLILTKKDFI +TLKYNNTLYPNFTIFSEIQKHWQPRIQMTKPIRPISYDEVCLNYLFSEEITNQVKLQTLA +KLNKEIHFNKSYHFNLKNRWLKQKLVINASTSKSTSSLLTNIHMDQGEDKKTSVGVSLKL +PAIYLCEEEGLHTNLFIHKAQRLLYKIKIILSEKALNVEHYSWNKNSSLQKTFGYQNGII +QAKSLLHTLPTNLNYNLKWINYKQKNIFTTNKVGFFFLKGNTFFNTSQKLFNKKITKQTT +FLNATIYNFGRNNKNNLISYNNSNFLENTHFVDLSLCLELQKIYLNKNYLNLKPILDKTI +HCQKPTKVLFKKSGFSKKQHYYLEFLNTKNHRRLIGLKEFNDYHMSYSKSQTKEMSNFID +SYYFVKPINMDCAHYIKHELVLYNDLITHFASLNLYISREHGLKSLSAFFINILKIFITS +NQSQISLAPIGIDKYTNIYIPEGEGEKDMTKNVFQVIKKSGQLIQMNKEKMTLRLGQPLV +ISPRSTIHATHGDFIRYKTPVVTLTYQQLKTGDIVQGIPKIEQLFEARTTKRGRLFRDNV +TNLLTGLFLKYFIKSTYLLRKTMIGFSKKRWKKSIKYTLPVNKQPNMPRVNHTLNTTVGT +ELGRQSKTKVDKNKHSIAINKNLNYSNFINNKQNQTIILALALQWAVKQSFYKIQQIIVD +GILRVYRSQGVSIADKHVEIVVKQMTSKVRIINSNASKMSEYMFSLDTIKAGEMPETDLP +EEEVSLQQNKAVSKQNVVAQTGKKRKKRLRKSKLSERDVITTKRTEGIDSSKIPSSNIPE +GKVTQNNKRKSTRKNVSLADRELKTRNTLSNTTKPIQISQVFEHKVLNQLLSNNLDGPTG +LFPGEIVDIDFVENINTFLLKTASVDRASRETLSTDPLNPNNQVAFAIEPIKYEPIVLGI +TRASLEVESFLSAASFQQTTRVLSQAALYKKKDFLKGLKENIIIGNLIPAGTGFLSSLNI +* +>sp|P19572| photosystem I P700 chlorophyll a apoprotein A2 GN=psaB PE=psaB.p01 +MATKLFPKFSQGLAQDPTTRRIWYGLAMAHDFESHDGMTEENLYQKIFASHFGQLSIIFL +WTSGNLFHVAWQGNFEQWVTDPVHIRPIAHAIWDPHFGQPAVEAFTRGGASGPVNISTSG +VYQWWYTIGMRTNQDLYVGSVFLALVSAIFLFAGWLHLQPNFQPSLSWFKDAESRLNHHL +SGLFGVSSLAWTGHLVHVAIPESRGQHVGWDNFLSVLPHPQGLTPFFTGNWAAYAQSPDT +ASHVFGTAQGSGQAILTFLGGFHPQTQSLWLTDMAHHHLAIAVIFIVAGHMYRTNFGIGH +RMQAILEAHTPPSGSLGAGHKGLFDTVNNSLHFQLGLALASVGTITSLVAQHMYSLPPYA +FQAIDFTTQAALYTHHQYIAGFIMCGAFAHGAIFFIRDYDPEQNKGNVLARMLDHKEALI +SHLSWVSLFLGFHTLGLYVHNDVMQAFGTPEKQILIEPVFAQWIQAAHGKALYGFDFLLS +SKTSAAFANGQSLWLPGWLDAINNNQNSLFLTIGPGDFLVHHAIALGLHTTTLILVKGAL +DARGSKLMPDKKDFGYSFPCDGPGRGGTCDISAYDAFYLAVFWMLNTIGWVTFYWHWKHL +TLWQGNVAQFDESSTYLMGWLRDYLWLNSSQLINGYNPFGMNSLSVWAWTFLFGHLIYAT +GFMFLISWRGYWQELIETLVWAHEKTPLANLVYWKDKPVALSIVQARLVGLAHFSVGYIF +TYAAFLIASTSGRFG* +>sp|P19573| ribulose-1,5-bisphosphate carboxylase/oxygenase large subunit GN=rbcL PE=rbcL.p01 +MVPQTETKAGAGFKAGVKDYRLTYYTPDYVVRDTDILAAFRMTPQPGVPPEECGAAVAAE +SSTGTWTTVWTDGLTSLDRYKGRCYDIEPVPGEDNQYIAYVAYPIDLFEEGSVTNMFTSI +VGNVFGFKALRALRLEDLRIPPAYVKTFVGPPHGIQVERDKLNKYGRGLLGCTIKPKLGL +SAKNYGRAVYECLRGGLDFTKDDENVNSQPFMRWRDRFLFVAEAIYKAQAETGEVKGHYL +NATAGTCEEMMKRAVCAKELGVPIIMHDYLTGGFTANTSLAIYCRDNGLLLHIHRAMHAV +IDRQRNHGIHFRVLAKALRMSGGDHLHSGTVVGKLEGEREVTLGFVDLMRDDYVEKDRSR +GIYFTQDWCSMPGVMPVASGGIHVWHMPALVEIFGDDACLQFGGGTLGHPWGNAPGAAAN +RVALEACTQARNEGRDLAREGGDVIRSACKWSPELAAACEVWKEIKFEFDTIDKL* +>sp|P19574| ATP synthase CF1 alpha subunit GN=atpA PE=atpA.p01 +MAMRTPEELSNLIKDLIEQYTPEVKMVDFGIVFQVGDGIARIYGLEKAMSGELLEFEDGT +LGIALNLEANNVGAVLLGDGLKITEGSRVRCTGKIAEIPVGEAYLGRVVDGLARPVDGKG +AVQTKDSRAIESPAPGIVARRSVYEPLATGLVAVDAMIPVGRGQRELIIGDRQTGKTAIA +VDTILNQKGKGVICVYVAIGQKASSVAQVLNTLKERGALDYTIIVMANANEPATLQYLAP +YTGATLAEYFMYTGRPTLTIYDDLSKQAQAYREMSLLLRRPPGREAYPGDVFYLHSRLLE +RAAKLNNALGEGSMTALPIVETQEGDVSAYIPTNVISITDGQIFLAAGLFNSGLRPAINV +GISVSRVGSAAQPKAMKQVAGKLKLELAQFAELEAFSQFASDLDQATQNQLARGARLREI +LKQPQSSPLSVEEQVASIYAGTNGYLDKLEVSQVRAYLSGLRSYLANSYPKYGEILRSTL +TFTPEAEGLVKQAINEYLEEFKSQAKAA* +>sp|P19575| photosystem II protein I GN=psbI PE=psbI.p01 +MLTLKIFVYTVVTFFVCLFIFGFLSNDPARNPGKNLD* +>sp|P19576| envelope membrane protein GN=cemA PE=cemA.p01 +MYTFIYFKNPKHFNLSQPSRCTAQPFISTRNRPLSCPIPTVQPIYVTHGIILPILKGTLI +SQRGVNIVPFVSKRDSSYYTDKDKVVSITYEEIGLFPRSFSRVLDRFLKQLFSDVDNLVI +QEYRFYRYLFLTTIKTIFILFFVPFLVNFAAKNYIVKPITEYFWNTSHPEIFLNSYEQKR +AFVELAKFEEKIYFETLVESHSHHQTHRDSKPLRENGIYFPDGEFLDNANLLSTPRSINS +NTFLKQNIDISLREEKPLTLVQGVNLLEEKKELNIPLAQENIAYNNQSIPQTSFGQGNFS +SLFTGDREGEETAKQNLLSQRVIGANLRQIYLPSAEGEMLPSIRGSLDSIKNKDISKIYQ +EKTIELATYYNNHSIEAITNFFADLLSLFTLLYLLITLEIQINITKSFLLEVFFGLDDSK +KSLLILLITDLLVGYHSSNLWELFFEFIFNHYGIPESQTGIFLLVATLPVLLDVLFKYLI +FRHLNRSSPATVATYQAIIE* +>sp|P19577| ATP synthase CF0 C subunit GN=atpH PE=atpH.p01 +MNPIVAAASVVSAGLAVGLAAIGPGMGQGTAAGYAVEGIARQPEAEGKIRGALLLSFAFM +ESLTIYGLVVALALLFANPFAG* +>sp|P19578| CF0 ATP synthase subunit I GN=atpF PE=atpF.p01 +MVFLPDNILILGHGGFGFNTNVFETNIINLAAVVGIVVSFVGKNLSSLLEDRKNTIVKNL +EEANQRAIEAEQKLTAARTQLETAKKKAQEIREEGVLRATQEINNVVSQHELRLARLQEF +KQETLAFYQQKAFKQAYLYVINKIMTRVRERLNKGLDSTYHVVVNNFYVSRFTQF* +>sp|P19579| ribosomal protein S11 GN=rps11 PE=rps11.p01 +MAKQTRKLTTNKTKKKIFRGVVHIQAGHHNTIVTITNIRGEVLCWSSAGACGFRGKRKST +SFAAKKAAETVARKSRDFTMKAAKILVTGPGQGRESAIREIFKAGIKVSVIREKTGIPHN +GCRPPKKRSV* +>sp|P19580| photochlorophyllide reductase subunit N GN=chlN PE=chlN.p01 +LLDGATTILNLNSFFECETGNYHTFCPISCVAWLYQKIEDSFFLVIGTKTCGYFLQNALG +VMIFAEPRYAMAELEESDISAQLNDYKELKRLCLQIKQDRNPSVIVWIGTCTTEIIKMDL +EGMAPRLETEIGIPIVVARANGLDYAFTQGEDTVLSAMALASLKKDVPFLVGNTGLTNNQ +LLLEKSTSSVNGTDGKELLKKSLVLFGSVPSTVTTQLTLELKKEGINVSGWLPSANYKDL +PTFNKDTLVCGINPFLSRTATTLMRRSKCTLICAPFPIGPDGTRVWIEKICGAFGINPSL +NPITGNTNLYDREQKIFNGLEDYLKLLRGKSVFFMGDNLLEISLARFLTRCGMIVYEIGI +PYLDKRFQAAELALLEQTCKEMNVPMPRIVEKPDNYYQIQRIRELKPDLTITGMAHANPL +EARGITTKWSVEFTFAQIHGFTNTREILELVTRPLRRNLMSNQSVNAIS* +>sp|P19581| ATP synthase CF1 beta subunit GN=atpB PE=atpB.p01 +MPWGILIPLTMSDSIETKNMGRIVQIIGPVLDIVFAKGQVPNIYNALTIRAKNSAGTEMA +VTCEVQQLLGDNCVRAVSMNPTEGLMRGMEVVDTGKPLSVPVGKVTLGRIFNVLGEPVDN +MGNVKVEETLPIHRTAPAFVDLDTRLSIFETGIKVVDLLAPYRRGGKIGLFGGAGVGKTV +LIMELINNIAKAHGGVSVFAGVGERTREGNDLYTEMKESGVIVEKNLSDSKVALVYGQMN +EPPGARMRVALTALTMAEYFRDVNKQDVLFFIDNIFRFVQAGAEVSALLGRMPSAVGYQP +TLATEMGGLQERITSTKDGSITSIQAVYVPADDLTDPAPATTFAHLDATTVLSRNLAAKG +IYPAVDPLESTSTMLQPWILGEKHYDSAQSVKKTLQRYKELQDIIAILGLDELSEEDRLI +VARARKIERFLSQPFFVAEVFTGSPGKYVSLAETIEGFGKIFAGELDDLPEQAFYLVGNI +TEAISKAASLK* +>sp|P19582| hypothetical protein GN=orf1995 PE=orf1995.p01 +MITFTFMSLVTSVKDYVEITHKLIEIEPLKNYTEFGAVFTYFIFSIGEFFKNFFSFSFLN +NIWSIPIIIPDIASAMISEVSVLDGYFHNAFTFLETSVNTTTNPSLVIFEKFVIGIINSL +FLILPTSTSHLITLRRFVMQGLEAGYMAGLGTLAGNFLWLASIILGWRFFVIPWLSLDIF +RYLLGFVLLVKYIWDSSKERRMALEDLSKWKIFLLNFLLALTEQSCIYPFISNLSFGPDA +SILEGFPVDNYPQFLLIHGAYLLGILFGSFSLLQFTCWFWENPAFSIYLWITTKSSLKIS +TSSYYKILNFTFLYATMLCAIASIPYYGLDYTITNPIGLVPQDRILNQKKSQSDPDKLIT +ETAFLNLNPTDKNSRIRDGVHARRERWKQRLIKYQAFDASTYDQGVYDFLTIEDLNYGFD +RFWLRRKMRNHQIRFRLFPGPWMRSLKKQLNNPANPSLETSTKAASGPRVEFFRILFEQF +YHPNFHDRAAMQTNPAEARNKFISTSPLASTESKKALNSTFSLGNINNSSTGIEGLVLTN +TQATLLPTDLQTKRTIKPGLIYTNSALRKFVRNVNTRLNVKLLNSKETNLTTKYKSQFIY +SKRWKSIFSKIQPLQNGTTRKSYQLFRNVAKQILVTPDAKSLKLITINQKLSLKERKLLE +LRTQYNNNSTLTTTAPLTLVRPLNVYLQKEEAFKRKLRYYGTMPMRKLTVGNQAPYFKAL +MKRGFYYYKPTLRWRKTLYVASLRRGFRKKSRKQRILVMPSNQQNFNNTLDNTKTNINQN +NLANPLGGNEVPMYGADGENSLITKPTHSYTVLGKRASRYRHQIYKDVLQHWYYTPFNRL +LMKFDVDAFINRQPKSHFLTKNEERALHIRRFLLSEHYDTLRWYTYMQHYKTMKTNIGGT +KSFANRAYNQQFQGTFKKIRHLFAITPKQGDFYTLKFDQPLYNDNKLKDNLYFHEELLTD +YYNGTNLQTNQTSNISVNSTTTFIDNSLRTTQLPVPSSSFDIVNQSSTLIGLTTMQNALR +KNVVESTLTSLNSDGEAATSQPKLNFVYSELFVKLIKECKKRIHDQTFLKNYITHRIEKR +EQLNQEQTKELNKRLEKLKVWLNSDKGSISKLQNTPVQDPNISSPDKVLTTAMQKAVNES +ISLSGIMPSDKIKTTYGNLTNAYTIKTENAILTKLNVINQLTNNETTTQKNTLIKSIGVN +KIQTVLQTIITNFKSSLYNQTQLLRVKTDKDLQWWRTKQRVITKRKSARKRDRFKKQIAV +VNKKLAALSKKVETEKSNLYQTLYGNYEISDYLLRNVPTGSSAVIDSTVLRKKQDNQAYL +PKETNNVQFNSFVDSNNNVWQTFFAKKLRKKISSKGRRYRSLSLARYLTATRKPRLVGLD +NLTKIDNITTLQGAFITKEEKQDSLNLTIQRKQELTNSLKKSQIKKRSRHSWKKRSRHQF +SRNHYKYRKRHTHGNGKLRVMNKKLKKFKATNELRQWWWNSFLPRYLSNLQVNNSTLTNK +NVSFKPLSNTNSVPSTNMASPTTSRNLLDNLNSSNQISTSASMNQNIVTESVKVETNQVY +LPEGEKSFDITSMTTTLPFYAGWDESLRKFVVTNRLLSRRDAGLSVNNNAQEINFTNAPI +QGLNEGSFLYWQTEMPFNSYNIDQFITTNQSFYAPLGWRRFEFRHSILKTWVNNTKAGNN +NIKKKTLIISLKNLQPLKSSQQKQNQIKTKKLVARRIKKRYKLLKQMPNQLMYSPTGPLL +TEVLPSHYISVFDQQYRLPRNRYLKRNPLKTLKKTTLLALMDSSKQTNGVNKEFTLRKRV +KPRRKYHRKRFIKKDGLIFPRRTKFNTNTTLTGNALITNNVNSIEEDDLRWRPSSRTKQK +RKDNTRSSAASKTKSNKRVKTNPLRLRQLRRREFQQVLKPLQRYIPQNGGFTWPGDYLRL +EIVEMPKLKSINIKKTSLKQKINVQPVGIMPRKYLIEKHNIKVLKKKLSQAYSTQQLTKV +VQEYKNLIQNSPPAI* +>sp|P19583| ribosomal protein S12 GN=rps12 PE=rps12.p01 +MPTIQQLIRSARKKITKKTKSPALKSCPQRRGICLRVYTVTPKKPNSALRKVARVRLTTG +FEVTAYIPGVGHNLQEHAVVLVRGGRVKDLPGVRYHIVRGSLDTAGVKNRVQSRSKYGVK +MGSKTAAKTAGKK* +>sp|P19584| photosystem I reaction center subunit IX GN=psaJ PE=psaJ.p01 +MKDFTTYLSTAPVIATIWFTFTAGLLIEINRYFPDPLVFSF* +>sp|P19585| ATP synthase CF0 A subunit GN=atpI PE=atpI.p01 +MNPLLEIAEVSVGQHYYWELGGYELHGQVLITSWVVLGILAVLSFLGNSNLKSTPDGFQN +FTELVTEFIRDLAKTQIGEEDYLKWVPFLGTIFLFIFVSNWSGALLPYRLIEIPNGELAA +PTNDINTTVALALLTSISYFYAGISKKGLGYFSRYVEPAAFLLPINVLEDFTKPLSLSFR +LFGNILADELVVGVLVALVPLIIPIPIMLLGVFTSAIQALVFATLAGAYIGEALADHH* +>sp|P19586| photosystem II reaction center protein J GN=psbJ PE=psbJ.p01 +MSNTGTTGRIPLWLVGTVVGTLAIGLLAVFFYGSYVGLGSSL* +>sp|P19587| photosystem II protein D2 GN=psbD PE=psbD.p01 +MTIAIGTYQEKRTWFDDADDWLRQDRFVFVGWSGLLLFPCAYFALGGWLTGTTFVTSWYT +HGLATSYLEGCNFLTAAVSTPANSMAHSLLFVWGPEAQGDFTRWCQLGGLWAFVALHGAF +GLIGFMLRQFEIARSVNLRPYNAIAFSAPIAVFVSVFLIYPLGQSGWFFAPSFGVAAIFR +FILFFQGFHNWTLNPFHMMGVAGVLGAALLCAIHGATVENTLFEDGDGANTFRAFNPTQA +EETYSMVTANRFWSQIFGVAFSNKRWLHFFMLLVPVTGLWMSAIGVVGLALNLRAYDFVS +QEIRAAEDPEFETFYTKNILLNEGIRAWMAAQDQPHERLVFPEEVLPRGNAL* +>sp|P19588| hypothetical protein GN=orf2971 PE=orf2971.p01 +MTFLNHYTYLFSIPEKQADKVSGILRLAQARPIETLQNERINKQLNAFLKTYKFEKLITN +YKKMQSFIPNNSLNGNKTNSSTNKLYATSLNVFPENPPLMVRKAVSDEADKFSKFTYSKV +QVVTNNLNNGMNSKEFIKANNLKPSLRAAESLVLNHLTYNKFKENLYFKTNNIQPTKSKS +TSLFFLNILSNSKPRTCSDFLSSPKIRKTWFRNTAWSLQTQQHRSSNGINLSLQLPYALG +PSVPAGASGQNMYELPVAQSSSRFGTYYFLQKLLSKYLDVWNASADNGSVLSNSENIKLN +FSMVSLLDSKMAIQTPNSLYFVFTQLNQKTFLSYWLLPVAGLALLTPTLLTLTGQSVSVQ +KFNSFINKKTDMMVLSNTEMPSKSFGTPTLFGTSVEIYLPNSYMPKGEGESGINRVNSSI +NAVKKNTVTANLVLDSESQEVATSFQNDLISIKYCFNNLYNYISNKTALSTKNLFLFSAI +KSNATKHKRTQSFFSVENTTTLGNNSNFVKGHFKSSINAFSSYLPSTNVHSMIPLTSLPY +LKAISPLYSKFMIDHSLKFITPKTTLKLLQHKLNKSPKQMYTKTQNFTGLRDLRALNSFS +FGQVNFRTNHFLHSNSRPLNHYNQALKLINGYEQYKNNLQINCNKTLDLNTKNKLVYQVN +KSHLFNQKCSQIVYKQSLYNRDLCTIRGTGTKVVDYFSHGDKLSNKNGIVLDYFVYSNLL +FDNKTNTIINKDGKQNITKLKLNLTKTTVPFKTLIKKYTSINSLVANEQTRNNLNLGLIH +FNGHLSVVSNANLLTGRPVKFIYYKFDKRLNSYLIYVNQNLKKFIQLNNNFLKPKPLSHQ +KNKPVEDFNQYATNNSSPPKTNVFEKSFVEDSSLRKPLTSLRGSKQFLNSLTILFKHQKM +FKKKTLKAHKWHSDTQGIFRKHTNSSFGSANFSNGPEESSLSTRLHIQKKRKAKKQRLET +RRQKKRTRFFPRPVWLRSRMFLNFLTERNKYYLNSTITKQGFSLPSKDVVTTKLDWLKED +MRRLPLGAYQYKSLLTQKAGNKFQRQSFTEVVSTMEYINGIHKALNNSIFNKIVRKSLLS +SSQNPLKLRLVANYSKMQFMHRVKLPFYRTLKHSEGTKNLANKKQNLRDIKIKANYNNFK +SQKANNQPQQNDKDKDKDTMFRDFWVWSYNNTQTNAFNQNLWWLLPNLTTKQSNLEFLTS +TYPTAKETQRAKEEIHGNSIPTASKNQIALIRLNWALNKTNINTFTDYSKRNNLWTTQKL +RNQSKNNKTKSLEKQFITNWEKFFLNKNLNIFSKKIISKVKQKKQKLNYMTSYLNVQSEH +NVKIFHNSWWTHLNIKNLVNNQDMVIPVREGYFSVGNFNSEFINSAIIKSINNKTLVENY +VYSPSSEKETMQLLLMSSSILLHLCAIISLVSISQVRCFVKFHLILLYKLSNVYNAILNQ +LSNKLQKNLPIYNNINKLNSRYFYMNHQKSQIKQRKKLLTYFSLTLLKKQFVTVKPLQIR +NFASIKNQSSNNSNLTYTDMLPLSLRANKFRGSKYDISIREEEGQSAHIKPSKSMYAKLN +ILSLKTIFLKQLLMNKKPSALPSNVGLKSNRETQKSQLIQRIKTKELQISLKKNIIGFSK +VTKNHILKILFNVIEVFQTAVRNISSFFEKPAEFTTTWIAYGFLVEWSSDFITIIPENVD +IYIWNVFSKIYRTIPLSFISTTLGPASTVFDPVTNSTIPIQMGNFNYQKMVAFPILLSLS +HLLHRRILYLFDTLFSTITQPDTDLIARQEKGTLFWDIWADFLVTAADYYNVNVAALSTI +KAEQNSLIENISNDFDNLTMSSKKPFFMPNKGVSNIKNIFWIKKLKEPQLPESIVQNREV +FVRERKRTLKGLFNIYAPQEETLWNNPTSPKNLSDEKISFKLFNQLNLQLFAEKNKIKPY +FEAYFSTTQQKTNIMQSAFPEANLNRWSVNQFITYQSWHSHNGSNNSNGDLFIDYHPPKT +FSHIPALKYNSILQQPIGSLVCQIYSGLFNKQISKNILLVNPKTTSNNLVDYNVLLIQAL +AGETEMKIITDNAQRYALVNRGFAIGIKLLREVFDAIALNTPCIFLLEDIHAIGERRPML +ISDFGGGMSDDNGSFKEDFFGSQRDEVHEKNQVVYQLTRHAITHYKKPFKGDYSLAIPTN +LYVTDLFLKLPTQSISNLTNVENHNLSIKNKIQHNGTQSLTETKRNLGGDINKNSYLQLT +QFTKTLAPPSTSPFSVLLLKEEKRLKPNKIVEELPWTSLPGEQLATKPRTSYSVRAKVAM +LAELSLSNLSAKLDMITDLLVIIDSVRSNKGFVVFATTDIPHVLDPALRRPGRLDETICL +PNIHTSNILNFTKNYEIFKSAKDTSNFGKKIILNEMQNLTTTSTQRDMYLSCLPTNNQTH +KTKREGVLTMNLKDYNILLNQVYFAEGTGGILNSQMHKDSLQKSLNFALISHSKKLKELN +VSKLIGSNGTVSQGNVDQLGVFAGQIVNKQKKSLQQHLPNSKKSFKKKYKDKAIIYYEVG +KFVLNYFLNNQLTQSSIIDKPVSVTNKQTNDITIFGNDFLNLKTINYLSLYNSKNKILLQ +LMLIFGGKISQLLSSKNLVKSLKQASINSYMVEEESGSISSAGMPLGQTHLLPKALSVLA +KPMIFSDGYNNQNLKTATTLLLSFIHKRYLYRKNLIVPKLLSFADGNILDEPPSPPFSSL +LIPAKRFENYKRFFRDTLTGDKMGQRKSQITLLEKLQYHMQLRSIKQLNATFSSQENLDF +QSNAALTSQKLDTLMSLSTNNLLQNPTNINWYYQNRILKRHGQYLTNQWWNGQLSEHNAE +TVFLSDIDWRSSFIKNKNINITKSKNLYRLTQQKNNTDGLDVLLDFPDTDQYYNPKRRRW +LLNNGSWNFWFNFDKLYSEEIVTTWILESLIQTYKYLHKNTELLDFVTNKFITLGYIAPE +NANLQNISGFPSQSELLSTKEIILTNSFKRF* +>sp|P19589| photosystem II 44 kDa protein GN=psbC PE=psbC.p01 +VETLFNGTLTVGGRDQETTGFAWWSGNARLINLSGKLLGAHVAHAGLIVFWAGAMNLFEV +SHFVPEKPMYEQGLILLPHIATLGYGVGPGGEIIDTFPYFVSGVLHLISSAVLGFGGVYH +SLIGPETLEESYPFFGYVWKDKNKMTNILGYHLIMLGLGAWLLVWKAMYFGGVYDTWAPG +GGDVRVITNPTTNAAVIFGYLVKSPFGGDGWICSVDNMEDIIGGHIWIGTLEILGGIWHI +YTTPWPWARRAFVWSGEAYLSYSLGAIGVMGFIACCMSWFNNTAYPSEFYGPTGPEASQS +QAFTFLVRDQRLGANVASAQGPTGLGKYLMRSPTGEIIFGGETMRFWDFRGPWLEPLRGP +NGLDLNKLKNDIQPWQERRAAEYMTHAPLGSLNSVGGVATEINAVNFVSPRSWLACSHFC +LGFFFFIGHLWHAGRARAAAAGFEKGIDRFDEPVLSMRPLD* +>sp|P19590| photosystem I subunit VII GN=psaC PE=psaC.p01 +MAHIVKIYDTCIGCTQCVRACPLDVLEMVPWDGCKASQMASAPRTEDCVGCKRCETACPT +DFLSVRVYLGSESTRSMGLSY* +>sp|P19591| cytochrome b6/f complex subunit VI GN=petL PE=petL.p01 +MLTITSYVGLLIGALVFTLGIYLGLLKVVKLI* +>sp|P19592| chloroplast RNA polymerase beta' chain GN=rpoC1b PE=rpoC1b.p01 +MSGQFTISDLNRLYQRIIYRNERLKKFLKDPALSSSFEMKYAQRLLQEAVDNLIQNGKSG +VVPEKDARGRLLKSLSDILKGKQGRFRQYLLGKRVDYSGRSVIVVGPRLRLHECGIPKEM +ALVLYSPFLIKRILNEKLADTYLSAKKLIRTNPLLVSQLLREIMKSCPVLLNRAPTLHRL +GFQAFQPKLVDGKAILLHPLVCPAFNADFDGDQMAVHVPITFEARAEAWKLMLARNNLLS +PATGEPLILPSQDMVLGCYYLTTNCAEKWSKLKKGSGMYFHNIHEVLKAYNQHLIHLHAV +IWVNIQGHLETANNIEQPLEIRIPLNQKYFYSQKGSYTALAGRPWQLLLREDAYSSNGLI +AEANAGNLPQINVAEGEVNKNILDLRYLNLNYMEIYSKTHNLLNVNHSTLVAQRGQIKDN +IHLPQLTTQIIRTTPGKILFNIIVKNAIEKRPKLLSKSNFKGGFIKNKLVNAIDQEIDNY +FIRKNTALS* +>sp|P19593| chloroplast RNA polymerase beta' chain GN=rpoC1a PE=rpoC1a.p01 +MHYKKYAELLKNGTYFSHIHSPTLLYPKGKVMYAQYGNSPLQGLNSQVTNLNLIRKVVAA +SAEMDSFKKLPNLQNNKLPLNNLKARLNTKAIYISNEEAQLTVSKIGTPNAIETDVPGTI +KNAPSNTLLTHRSFKFFVNKIYAKPLISLKKTKETKPLLLGKTGTTLTQKGLNPFQSLFL +NTKSSSPSTARSFGTKNIVNTLQIKKIVHKFENSYSKLTEINLITINLASANRIRQWAEK +TLPNGKVVGEVINPETIHYKTLKPIKGGLFCERIFGPLKDHECACGKKFNIKNYLTKTVN +NSSIQPIAESRQTQPNTQLSLNLQKRYFCRICDVEYTYSIIRRTQLGYIQLASPTTHVWF +VKGIPSYISILLDMKKKHLQGITYNTETLTLENSFRGRQLLPVSPSSIFESWQKIMKKQY +PEKYNLTNTMIKIKSTNTLPLRVSQPNSNTSYSYMPNIYIPEGEGEEKTKTKLKKTTPLN +AIAQKGVKYKKPKTKKALYKQYLKNYYARKPKNSNQAEAVSFGVNKVQLPLTAFQQRNKV +YFYKSEKRNWPVLTTVAKMQYIVSKKGWFKLIQYVIKSAESYGAADSI* +>sp|P19594| cytochrome b GN=cob PE=cob.p01 +MRMHNKIQLLSVLNTHLVAYPTPMNLNYSWNGGSLAGMMLASQMLTGILLAMHYVGHVDY +AFASVQHLMTDVPSGMILRYAHANGASLFFIVVYLHVLRGMYYGSGAQPREIVWISGVVI +LLVMIITAFIGYVLPWGQMSFWGATVITSLATAIPVVGKHIMYWLWGGFSVDNPTLNRFY +SFHYTLPLILAGLSVFHIAALHQYGSTNPLGVNSQSSLISFGSYFGAKDLVGALFLALVF +SILVFFYPDLLGHPDNLIPANPYSTPQHIVPEWYFLWVYAILRSIPNKAMGVLAIGLVFA +SLFAMPFIGLGGGKFRIITEWLYWTFLADVLLLTWLGGNEITPITSFVGQCCTAYLFFYL +LVCQPLVGYLETQFAHGTQTN* +>sp|P19595| NADH dehydrogenase subunit 4 GN=nad4 PE=nad4.p01 +MFISVLLILFALCVTLIPEAHYHMVRVWSFVATIIPMWVVTWMWWNFDASGHGLQMLVIL +GRSHLAFGIDGVALSLMLLTTVLFPICMMLLRTVAGFMTFILLEVLVLSALCVLDLLGFY +ILFEASLILLFLLIGRAPYGSLEAAYKIVLYTMAGSLVLLPTLFMIYSECGTTNVLYMTC +AYNHQTVLGWGLLAVLAVKIPLMPVHLWLPEAHVAAPTAGSVLLAGVLLKLGGIGFLRFM +LPVVPEFCVSVFPLVSTLCLVSFLFSTLSTLRQIDLKKIVAYSSIAHMSMVTLAIFSQSE +FSAYSSSFLMIAHGLISPALFLIVGILYDRAHTKFILYFSGLGASMPIGSTLFFLFTLGN +LAFPLFPNFIAEVLCMVSIFAVHELLAYVFCVCQVLGAAYGFWAFNRVVHGLPRGPADVT +RTEFHTVLPLLIGAVWLGIKPMA* +>sp|P19596| NADH dehydrogenase subunit 5 GN=nad5 PE=nad5.p01 +MFLLSVFFPLLGGLVNTSPIARFLGHRGSSIIAIGCMVVAFISSVVIYYEVVFMGCAVSV +DVFGTWFSVGTFHAGWTFNFDLLTANMLFTVTGVSMAVHMYACDYMRQDPHLNLFLGYLS +YFTGFMCVLVAADNLLVMLVGWEGIGVCSYLLIGYWSHRLSAVKSAQKAILVNRVSDGLL +MWGVLWVWYHLGSLEYDLLNVYSASGFVGLSILIGAMGKSAQILFHVWLADAMEGPTPVS +ALIHAATLVTAGVYLLVRLHIHDEMFVIIVGSLTAFMAGVFGATQSDLKRVIAYSTCSQL +GYMMVSLGLGETGGEASMGHLMTHASFKAALFLAAGMVISGNGGNQHIARYGGSAHSAMF +TMLTLMVASLSLIGWPELSGFYSKETILNLAAICADPIADVAHTLLLLTAMLTSAYTTKL +FYQCFMVDFSGSSVTPVRNVLPILAMAILLLDIMLKVWVGTNLLSGMLFFLPWGVKTLPF +GLMVAGILTATAAVGSERFTLIRFCGSRWGFDQLFARSPVNPIFDLGRITWAIGDRGLLS +VGNLRA* +>sp|P19597| cytochrome oxidase subunit 1 GN=cox1 PE=cox1.p01 +MRWLYSTSHKDIGLLYLVFAFFGGLLGTSLSMLIRYELALPGRGLLDGNGQLYNVIITGH +GIIMLLFMVMPALFGGFGNWLLPIMIGAPDMAFPRLNNISFWLNPPALALLLLSTLVEQG +PGTGWTAYPPLSVQHSGTSVDLAILSLHLNGLSSILGAVNMLVTVAGLRAPGMKLLHMPL +FVWAIALTAVLVILAVPVLAAALVMLLTDRNINTAYFCESGDLILYQHLFWFFGHPEVYI +LILPAFGIVSQVVSFFSQKPVFGLTGMICAMGAISLLGFIVWAHHMFTVGLDLDTVAYFT +SATMIIAVPTGMKIFSWMATIYSGRVWFTTPMWFAVGFICLFTLGGVTGVVLANAGVDML +VHDTYYVVAHFHYVLSMGAVFGIFAGVYFWGNLITGLGYHEGRAMVHFWLLFIGVNLTFF +PQHFLGLAGMPRRMFDYADCFAGWNAVSSFGASISFISVIVFATTFQEAVRTVPRTATTL +EWVLLATPAHHALSQVPVLRTASSH* +>sp|P19598| NADH dehydrogenase subunit 2 GN=nad2 PE=nad2.p01 +MIELDLCFGLLLLILFGLLSLRNGHVSLAHIRFICQCWLVMITPLEVQDFALCILTVVLL +QSFHSFEALLFLLLAYIGQLYMMHSCNLVSFYVCLEAQTLCVVVLCGLLARGASTSFSVE +AALKFLLLSAMVSGMALFWFSAMYQRTGSLDMVGQETFWILLVMLFKLGVAPMHMWSVDL +YGSIPKSLLLYLSTAPKLSLFTFWASSWHHDFSVGVFILFSMFIGSIGAYGQPALRSLFA +YSTINEIGLLLLAVETAGFHTLYQHLGIYIITQLLLWNLTDKRLFALCAVSLAGLPPFAG +FFGKAWIFWHAMSVQAFSLLAAALFCTLLSLVYYLRVIRLFWTAPVHTAASFTGAPNQTT +LTSACAVALAFAPVMLVKPFVI* +>sp|P19599| NADH dehydrogenase subunit 6 GN=nad6 PE=nad6.p01 +MFFENSAILLCALLSIAVGYTKSPFMSLMYSVMLFINSSFVLMMLGFEFLALVNLLVYVG +ALAVLFLFVIMLLEIPATELRAYSRGWSTLGIFVFIINGVFQITPSMGPRGIITGLPGAE +SITNLGHALYLYFADLLILNSLVLTVALFGRFAIAPVRTTGR* +>sp|P19600| NADH dehydrogenase subunit 1 GN=nad1 PE=nad1.p01 +MIVASILILIVPVLLSVAMFTLAERTVMASMQRRFGPQVSGISGLLQPFWDGLKLGVKEP +VLPDSSSAGAFAASPMISFVLSQVAWVGICISDASFQGLVIMAISSLAVYGVMLAGWASN +SKYAFLGCLRSVALMVSYELSLGAALLSIGLFVTDGTGMKCLNFAEMPTTPQYAMLPLCL +IFLVCILAETKRDPFDLPEAELVAGYNVEYSSLGFALFFIAEYANMAVMSAIASIYFLGG +FSALKITALFFAFVWTRGTLPRYRYDQFMRLGWKAFLPLTLAFFALHASVAI* +>sp|P19601| reverse transcriptase-like protein GN=rtl PE=rtl.p01 +MFTVNYILDLFHTINATEVQHTLMHKRPASLHEVLRLAGLLQGYKDFLSLTKQAEAFFQA +GDYRSAFNTISKMWVHPQIVPALLLMECFETRELTVPYWYKSIKLRRQYIQKKNGGVRPI +IVAHKGLRICMAVINRLLQSCCVSWSNQTFGFRPGFGTHHAVLHLAQKAQQMLNKQQQAV +LVTFDLQAAYNSVDIKHLMQTLQLQYLPNDMKKLIWMWQHLPLAQLNAGINGLAQGYAYS +PTLFAWYVDQLVGQHMDFTIYADNFAGVFLTQQDAQFAVKEAQTLLQKSGLLIAPSSIKM +HLLDKNQHSELNWLGHKVLFPSCTVKLQHHQLLVNQHSPQVWTIQKWDKMLRTLGWVQLA +LNADWRRF* \ No newline at end of file From 91f948619b8baf5006ff853101757ea2bcde5aba Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 22:01:57 +0200 Subject: [PATCH 07/11] Add SEQUEST-like scoring documentation --- docs/02_03_sequest_like_scoring.fsx | 402 ++++++++++++++++++++++++++++ 1 file changed, 402 insertions(+) create mode 100644 docs/02_03_sequest_like_scoring.fsx diff --git a/docs/02_03_sequest_like_scoring.fsx b/docs/02_03_sequest_like_scoring.fsx new file mode 100644 index 0000000..a5ff05a --- /dev/null +++ b/docs/02_03_sequest_like_scoring.fsx @@ -0,0 +1,402 @@ +(** +--- +title: SEQUEST-like scoring +category: Peptide identification +categoryindex: 2 +index: 3 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# SEQUEST-like scoring + +Identifying the peptide behind an MS2 spectrum runs through five steps: the +measured spectrum is preprocessed ([signal detection]({{root}}01_02_signal_detection.html)), +candidate sequences are selected by precursor mass +([peptide search databases]({{root}}02_02_search_databases.html)), a theoretical +spectrum is predicted for every candidate, the predictions are matched against the +measurement, and the closeness of fit is scored. This page walks steps three to five +for one spectrum, using the `SequestLike` module. + +The scoring question sounds simple: which candidate's predicted peaks line up +best with the measured ones? Representing both sides as intensity vectors +over 1 Da bins makes a dot product the obvious measure of agreement. On its +own, that measure has a flaw. A measured spectrum with many peaks all over +the m/z axis gets a decent dot product with almost any candidate, simply +because most predictions land near some peak. SEQUEST's cross-correlation +score (xcorr) fixes this by asking how much better the prediction correlates +with the measured spectrum at zero offset than on average across a range of +shifted offsets. That mean off-position correlation is the background, and +subtracting it leaves only the agreement that is specific to the candidate's +peak pattern. + +The `SequestLike` implementation follows this idea in a form that lets one +spectrum be scored against many candidates cheaply. The measured spectrum is +binned to 1 Da, and its intensities are square-root scaled and normalized to +the maximum within each of 10 windows. The average of the spectrum's shifted +copies over a delay range of 75 is then subtracted once. Algebraically that +folds the whole background subtraction into the measured side, so scoring a +candidate afterwards reduces to a dot product between the candidate's +predicted intensity vector and this preprocessed measured vector. The module is deliberately named SEQUEST-like. +Its background window differs from the symmetric average in the published +SEQUEST algorithm, so scores follow the same idea as the original without +matching its numbers exactly, and they are comparable within this +implementation. + +## Loading the measured spectrum + +The running example of these pages is the MS2 scan of the doubly charged +peptide ANLGMEVMHER in `ms2Example.mgf`. Its precursor m/z of 643.803548 and +charge 2 were recovered on the +[charge state determination]({{root}}01_03_charge_state_determination.html) +page. `PeakArray.zip` turns the two raw arrays of the reader into the +`PeakArray` the scorer consumes, as introduced on the +[peaks and peak arrays]({{root}}01_01_peaks_and_peak_arrays.html) page. +*) + +open BioFSharp +open BioFSharp.FileFormats.MGF +open BioFSharp.IO +open BioFSharp.Mz + +let ms2 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms2Example.mgf") + |> List.head + +let precursorMZ = + match MGFEntry.tryGetPrecursorMZ ms2 with + | Some mz -> mz + | None -> failwith "no precursor m/z in the MS2 header" + +let spectrum : PeakArray = PeakArray.zip ms2.Mass ms2.Intensity + +printfn "peaks: %i covering m/z %.2f to %.2f" + spectrum.Length spectrum.[0].Mz spectrum.[spectrum.Length - 1].Mz + +(** +```text +peaks: 971 covering m/z 100.67 to 1337.63 +``` + +## Assembling the candidate list + +In a production pipeline the candidates come out of a +[search database]({{root}}02_02_search_databases.html) query for all peptides +within a narrow mass window around the measured precursor mass. To keep this +page self-contained we build such a result list by hand with +`SearchDB.createLookUpResult`, which produces the same `LookUpResult` records +the database returns. The neutral mass of each candidate is computed in code +as its residue masses summed up plus one water for the termini. + +The list holds the true peptide and three pretenders. Two are permutations +of ANLGMEVMHER that keep the C-terminal arginine, so they weigh exactly the +same. The third replaces the asparagine with two glycines, a substitution +that is also exactly isobaric because the asparagine residue and two glycine +residues share the elemental composition. A mass window query cannot +distinguish any of these by mass. Only the fragment pattern can. +*) + +let mono : IBioItem -> float = BioItem.monoisoMass + +let neutralMass (peptide: AminoAcids.AminoAcid list) = + (peptide |> List.sumBy mono) + mono ModificationInfo.Table.H2O + +// Stand-ins for what a search database mass window query would return; +// built by hand to keep the page self-contained. +let candidate modSeqId pepSeqId (sequence: string) = + let bioSequence = BioList.ofAminoAcidString sequence + let mass = neutralMass bioSequence + let roundedMass = int64 (System.Math.Round(mass * 1000000.)) + SearchDB.createLookUpResult modSeqId pepSeqId mass roundedMass sequence bioSequence 0 + +let candidates = + [ candidate 1 1 "ANLGMEVMHER" // the peptide the scan was recorded from + candidate 2 2 "MANGLEVMHER" // permutation, same mass + candidate 3 3 "EVMANLGMHER" // permutation, same mass + candidate 4 4 "AGGLGMEVMHER" // Asn replaced by Gly-Gly, also isobaric + ] + +candidates +|> List.iter (fun c -> printfn "%-13s %10.4f Da" c.StringSequence c.Mass) + +(** +```text +ANLGMEVMHER 1285.5907 Da +MANGLEVMHER 1285.5907 Da +EVMANLGMHER 1285.5907 Da +AGGLGMEVMHER 1285.5907 Da +``` + +All four candidates sit at 1285.59 Da, within about two millidaltons of +the neutral precursor mass determined from the MS1 scan. + +## Predicting a theoretical spectrum per candidate + +Each candidate is paired with its predicted fragment masses. +`Fragmentation.Series.fragmentMasses` produces them exactly as on the +[in silico fragmentation]({{root}}02_01_fragmentation.html) page: the b and y +series of the candidate sequence as `TargetMasses`, and the same series of +the reversed sequence as `DecoyMasses`. We use monoisotopic masses +throughout. + +`SequestLike.getTheoSpecs` then converts every pair into a +`TheoreticalSpectrum`, holding one binned intensity vector for the target and +one for the decoy. The intensity of each predicted peak comes from a simple +model (`predictIntensitySimpleModel`): main series ions get the full +predicted intensity, loss peaks and minor series a fraction of it, +everything divided by the charge the ion is predicted at. Every fragment is +laid down once per charge from 1 up to the precursor charge, and the vector +is binned to 1 Da like the measured side, so the two can be compared bin by +bin. + +The scan limits define the m/z range of that binning. We use 100 to 1300, +which covers the recorded peaks starting at m/z 100.67 as well as the +heaviest interesting fragment, the singly protonated full-length y ion at +m/z 1286.6. The upper border only cuts away a single stray peak at m/z +1337.6, and no peak of the scan rounds exactly onto a border, where the +binning would drop it. +*) + +let scanlimits = 100., 1300. + +let fragmentPairs = + candidates + |> List.map (fun c -> + let fragments = + Fragmentation.Series.fragmentMasses + Fragmentation.Series.bOfBioList + Fragmentation.Series.yOfBioList + mono + c.BioSequence + c, fragments) + +let theoSpecs = SequestLike.getTheoSpecs scanlimits 2 fragmentPairs + +open FSharp.Stats + +let countOccupied v = + v |> Vector.toArray |> Array.filter (fun x -> x > 0.) |> Array.length + +printfn "bins per vector: %i" (Vector.length theoSpecs.Head.TheoSpec) + +theoSpecs +|> List.iter (fun ts -> + printfn "%-13s target bins occupied: %i decoy bins occupied: %i" + ts.LookUpResult.StringSequence + (countOccupied ts.TheoSpec) + (countOccupied ts.DecoyTheoSpec)) + +(** +```text +bins per vector: 1200 +AGGLGMEVMHER target bins occupied: 90 decoy bins occupied: 90 +EVMANLGMHER target bins occupied: 104 decoy bins occupied: 105 +MANGLEVMHER target bins occupied: 97 decoy bins occupied: 96 +ANLGMEVMHER target bins occupied: 97 decoy bins occupied: 98 +``` + +Every candidate now owns two binned prediction vectors, each occupying +around a hundred of the 1200 bins. `getTheoSpecs` builds its result list by +prepending, so the order is reversed relative to the input, which does not +matter because the scorer ranks by score anyway. + +## Preprocessing the measured spectrum + +The scorer performs the background subtraction on the measured side once, through +`spectrumToIntensityArrayMinusAutoCorrelation`. `calcSequestScore` calls it +internally, so this block is purely illustrative. +*) + +let preprocessed = + SequestLike.spectrumToIntensityArrayMinusAutoCorrelation scanlimits spectrum + +let values = preprocessed |> Vector.toArray + +printfn "vector length: %i" values.Length +printfn "positive entries: %i" (values |> Array.filter (fun x -> x > 0.) |> Array.length) +printfn "negative entries: %i" (values |> Array.filter (fun x -> x < 0.) |> Array.length) + +(** +```text +vector length: 1200 +positive entries: 407 +negative entries: 793 +``` + +The negative entries are the signature of the background subtraction. A bin +whose intensity is lower than the local self-correlation average now +penalizes a candidate that predicts a peak there, while a bin that stands +out above the background rewards it. The dot product with a prediction +vector therefore measures alignment beyond what shifted versions of the +spectrum would produce by chance. + +## Scoring the candidates + +`SequestLike.calcSequestScore` takes the scan limits, the measured spectrum, +the scan time, the precursor charge, the isolation window target m/z (the +precursor m/z from the header), the theoretical spectra and a free-text +spectrum identifier. It scores every target and every decoy vector against +the preprocessed measured vector and returns one `SearchEngineResult` per +scored spectrum, ranked by descending score. +*) + +let results = + SequestLike.calcSequestScore + scanlimits spectrum 20.93 2 precursorMZ theoSpecs "ms2Example" + +let printRanked (rs: SearchEngineResult.SearchEngineResult list) = + printfn "%-13s %-6s %8s %12s %8s" "sequence" "target" "score" "dBestToRest" "dNext" + rs + |> List.iter (fun r -> + printfn "%-13s %-6b %8.4f %12.4f %8.4f" + r.StringSequence r.IsTarget r.Score r.NormDeltaBestToRest r.NormDeltaNext) + +printRanked results + +(** +```text +sequence target score dBestToRest dNext +ANLGMEVMHER true 13.0383 0.0000 0.0132 +AGGLGMEVMHER true 12.8665 0.0132 0.3777 +MANGLEVMHER true 7.9418 0.3909 0.2979 +EVMANLGMHER true 4.0572 0.6888 0.1270 +ANLGMEVMHER false 2.4008 0.8159 0.0282 +AGGLGMEVMHER false 2.0329 0.8441 0.1274 +MANGLEVMHER false 0.3723 0.9714 0.0244 +EVMANLGMHER false 0.0548 0.9958 0.0000 +``` + +The target of ANLGMEVMHER comes out on top at 13.04. The two permutations +score far below it, at 7.94 and 4.06. Both end in the same MHER stretch and therefore +predict the true peptide's low y ions, which is why they still beat every +decoy, while their remaining predictions land in the wrong bins. +MANGLEVMHER even shares the y ladder up to y6 with the true sequence, and +that larger overlap is its lead over the other permutation. + +The Gly-Gly candidate almost ties the true peptide at 12.87: two glycines weigh +exactly as much as one asparagine, so from b3 onward its b ladder reproduces the +true one bin for bin, and the shared C-terminal nine residues make y1 to y9 +identical as well. Only a handful of bins differ, among them the extra b2 of the +glycine pair, and those few bins are the entire margin. Under any mass-based +fragment comparison a sequence whose fragments are isobaric with the true ones is +close to indistinguishable, and the small `dNext` of the top hit records that +ambiguity. + +## Reading a SearchEngineResult + +Printing the best hit in full shows everything the record carries. +*) + +printfn "%A" results.Head + +(** +```text +{ SearchEngine = SEQUESTLike + SpectrumID = "ms2Example" + ModSequenceID = 1 + PepSequenceID = 1 + GlobalMod = 0 + IsTarget = true + ScanTime = 20.93 + StringSequence = "ANLGMEVMHER" + PrecursorCharge = 2 + PrecursorMZ = 643.8035478 + MeasuredMass = 1285.592543 + TheoMass = 1285.590726 + PeptideLength = 11 + Score = 13.03831064 + NormDeltaBestToRest = 0.0 + NormDeltaNext = 0.01317481341 } +``` + +`SearchEngine` names the scorer that produced the record, so results of +different engines can share one result type. `SpectrumID` is the identifier +string passed into the call, and `ScanTime` travels along the same way, so a +PSM can be traced back to its scan. `ModSequenceID` and `PepSequenceID` are +copied from the `LookUpResult` and tie the hit back to the +[search database]({{root}}02_02_search_databases.html) rows it came from, +and the `GlobalMod` labeling flag rides along with them. +`IsTarget` distinguishes the candidate's own fragment prediction from that +of its reversed decoy sequence. Both records of such a pair share all +identifiers, since they stem from the same database entry. + +`StringSequence`, `PrecursorCharge`, `PrecursorMZ` and `PeptideLength` +describe the match itself. `MeasuredMass` is the neutral mass computed from +the isolation window target m/z and the charge, while `TheoMass` is the +candidate's database mass, so the difference between the two is the +precursor mass error of the match. `Score` is the xcorr-style dot product. + +The two delta fields put each score into the context of the whole ranking. +`calcNormDeltaBestToRest` fills `NormDeltaBestToRest` with (best score minus this +score) divided by the best score, so the best hit gets 0 and weaker hits approach 1: +MANGLEVMHER's 0.39 means it lost 39 percent of the top score. `calcNormDeltaNext` +fills `NormDeltaNext` with the gap to the next-ranked PSM, normalized by the best +score, with the last PSM getting 0: the top hit's 0.0132 is the Gly-Gly near-tie +discussed above. Both functions expect their input ranked by descending score, which +`calcSequestScore` arranges internally, and a best score of zero or below makes them +return sentinel values instead, 1 and 0 respectively for every PSM. How far a hit +stands out from its competitors is an input to +[false discovery rate control]({{root}}04_01_fdr_control.html). + +## What the decoys are for + +The ranked table contains eight PSMs for four candidates because every candidate +was also scored as its reversed decoy, the pairing introduced on the +[in silico fragmentation]({{root}}02_01_fragmentation.html) page. All four decoy +scores sit between 0.05 and 2.40, the level a wrong sequence of the right mass +reaches on this spectrum. A single xcorr value has no absolute meaning, and only the +comparison against a population of known wrong matches tells whether a 13 is +convincing. Collected over a whole run, the decoy scores estimate the score +distribution of chance matches, which is what FDR control is built on. + +## Scoring many spectra + +For a whole run with thousands of spectra, `calcSequestScoreParallel` +distributes the per-candidate scoring with `Async.Parallel`. It takes the +same arguments and produces the same ranking. +*) + +let resultsParallel = + SequestLike.calcSequestScoreParallel + scanlimits spectrum 20.93 2 precursorMZ theoSpecs "ms2Example" + +let bestSequential = results.Head +let bestParallel = resultsParallel.Head + +printfn "sequential best: %-13s target=%b score=%.4f" + bestSequential.StringSequence bestSequential.IsTarget bestSequential.Score +printfn "parallel best: %-13s target=%b score=%.4f" + bestParallel.StringSequence bestParallel.IsTarget bestParallel.Score + +(** +```text +sequential best: ANLGMEVMHER target=true score=13.0383 +parallel best: ANLGMEVMHER target=true score=13.0383 +``` + +## Where the scores go next + +The library offers further scorers following a different scoring philosophy, +covered in +[Andromeda-like and X!Tandem-like scoring]({{root}}02_04_andromeda_like_scoring.html), +and the target and decoy scores collected across a run feed +[false discovery rate control]({{root}}04_01_fdr_control.html). +*) From 323dea79f377e91ab1a6625e2a30ff388981f62b Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 22:13:16 +0200 Subject: [PATCH 08/11] Add Andromeda-like and X!Tandem-like scoring documentation --- docs/02_04_andromeda_like_scoring.fsx | 336 ++++++++++++++++++++++++++ 1 file changed, 336 insertions(+) create mode 100644 docs/02_04_andromeda_like_scoring.fsx diff --git a/docs/02_04_andromeda_like_scoring.fsx b/docs/02_04_andromeda_like_scoring.fsx new file mode 100644 index 0000000..315eb24 --- /dev/null +++ b/docs/02_04_andromeda_like_scoring.fsx @@ -0,0 +1,336 @@ +(** +--- +title: Andromeda-like and X!Tandem-like scoring +category: Peptide identification +categoryindex: 2 +index: 4 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Andromeda-like and X!Tandem-like scoring + +The [SEQUEST-like scoring]({{root}}02_03_sequest_like_scoring.html) page +turned the agreement between a predicted and a measured spectrum into a dot +product of intensity vectors. The scorers on this page, `AndromedaLike` and +`XScoring`, ask a probability question. +Given that a candidate predicts n fragment peaks inside the measured range +and k of them coincide with a measured peak, how surprising is that under +random matching? The answer is a cumulative binomial probability, the chance +of seeing at least k hits among n tries when each try succeeds with a fixed +background probability. The score reports that probability as a -10 log10 +value, so a higher score means the match is less likely to be chance. This +is the scoring idea of Andromeda, the search engine behind MaxQuant, and the +same matching machinery also yields an X!Tandem-style hyperscore. + +Intensity enters through which measured peaks are allowed to match at all, +an idea called peak depth. `ratedSpectrum` rates every measured peak by +counting how many more intense peaks lie within a 100 Da window centered on +it. At depth q, only peaks that rank among the q most intense within their +own window are offered to the matcher, and q also sets the background +probability of a random hit to q/100 (capped at 0.5 in the implementation), +one window of width 100 Da holding q acceptable peaks. `countMatches` tallies n and k for every depth of a +user-given range in a single pass over the predictions. `scoreFuncImpl` then +turns each (n, k, q) triple into a score, and the best score across the +depths is kept. The trade-off behind the range: a small q admits only the +strongest peaks, so a random match is improbable, but genuine fragments of +modest intensity are missed. A large q admits more real fragments along with +more noise. Evaluating a range of depths lets every candidate be scored at +the depth that suits the spectrum best. + +## Loading the measured spectrum + +The measured side is the running example of these pages, the MS2 scan of the +doubly charged peptide ANLGMEVMHER from `ms2Example.mgf` with its precursor +at m/z 643.803548. The scorer locates matching peaks with a binary search, +so the measured spectrum must be sorted ascending by m/z, which this scan +is. +*) + +open BioFSharp +open BioFSharp.FileFormats.MGF +open BioFSharp.IO +open BioFSharp.Mz + +let ms2 = + MGF.read (__SOURCE_DIRECTORY__ + "/data/ms2Example.mgf") + |> List.head + +let precursorMZ = + match MGFEntry.tryGetPrecursorMZ ms2 with + | Some mz -> mz + | None -> failwith "no precursor m/z in the MS2 header" + +let spectrum : PeakArray = PeakArray.zip ms2.Mass ms2.Intensity + +printfn "peaks: %i covering m/z %.2f to %.2f" + spectrum.Length spectrum.[0].Mz spectrum.[spectrum.Length - 1].Mz + +(** +```text +peaks: 971 covering m/z 100.67 to 1337.63 +``` + +## Rebuilding the candidate list + +The candidates are the same four isobaric sequences the +[SEQUEST-like page]({{root}}02_03_sequest_like_scoring.html) built and discussed in +detail. All four weigh 1285.5907 Da, so a mass window query cannot tell them apart. +*) + +let mono : IBioItem -> float = BioItem.monoisoMass + +let neutralMass (peptide: AminoAcids.AminoAcid list) = + (peptide |> List.sumBy mono) + mono ModificationInfo.Table.H2O + +// Stand-ins for what a search database mass window query would return; +// built by hand to keep the page self-contained. +let candidate modSeqId pepSeqId (sequence: string) = + let bioSequence = BioList.ofAminoAcidString sequence + let mass = neutralMass bioSequence + let roundedMass = int64 (System.Math.Round(mass * 1000000.)) + SearchDB.createLookUpResult modSeqId pepSeqId mass roundedMass sequence bioSequence 0 + +let candidates = + [ candidate 1 1 "ANLGMEVMHER" // the peptide the scan was recorded from + candidate 2 2 "MANGLEVMHER" // permutation, same mass + candidate 3 3 "EVMANLGMHER" // permutation, same mass + candidate 4 4 "AGGLGMEVMHER" // Asn replaced by Gly-Gly, also isobaric + ] + +printfn "%s" (candidates |> List.map (fun c -> c.StringSequence) |> String.concat ", ") + +(** +```text +ANLGMEVMHER, MANGLEVMHER, EVMANLGMHER, AGGLGMEVMHER +``` + +## Predicting tagged fragment spectra + +The fragment masses per candidate come from +`Fragmentation.Series.fragmentMasses` with the b and y series, exactly as on +the SEQUEST-like page. `AndromedaLike.getTheoSpecs` then converts every pair +into a `TheoreticalSpectrum` whose target and decoy sides are arrays of +`PeakFamily`, predicted m/z values that keep their ion series +tags. The prediction model of this scorer family: a singly charged precursor +gets singly charged predicted fragments only, and a precursor of charge two +or more gets every fragment as a singly and a doubly charged predicted peak +(higher fragment charges are not emitted). The neutral-loss dependents ride +on the singly charged copies. The peaks carry no predicted +intensities, matching needs only the m/z and the tag. Restricting the +comparison to the scan limits happens in the matching step, which skips +predicted peaks outside the range. +*) + +let scanlimits = 100., 1300. + +let fragmentPairs = + candidates + |> List.map (fun c -> + let fragments = + Fragmentation.Series.fragmentMasses + Fragmentation.Series.bOfBioList + Fragmentation.Series.yOfBioList + mono + c.BioSequence + c, fragments) + +let theoSpecs = AndromedaLike.getTheoSpecs scanlimits 2 fragmentPairs + +theoSpecs +|> List.iter (fun ts -> + let withLosses = + ts.TheoSpec + |> Array.filter (fun f -> not f.DependentPeaks.IsEmpty) + |> Array.length + printfn "%-13s peak families: %i with loss dependents: %i" + ts.LookUpResult.StringSequence ts.TheoSpec.Length withLosses) + +(** +```text +AGGLGMEVMHER peak families: 48 with loss dependents: 18 +EVMANLGMHER peak families: 44 with loss dependents: 22 +MANGLEVMHER peak families: 44 with loss dependents: 20 +ANLGMEVMHER peak families: 44 with loss dependents: 21 +``` + +The 22 b and y fragments of an 11 residue candidate become 44 peak families, +one per charge state, and the extra residue of the Gly-Gly variant adds two +more fragments and four more families. The families that carry loss +dependents are the singly charged copies of fragments containing loss-prone +residues. + +## Scoring at increasing peak depths + +`AndromedaLike.calcAndromedaScore` takes the depth range, the scan limits, +the matching tolerance in ppm of the fragment m/z, the measured spectrum, +the scan time, the precursor charge and isolation window target m/z, the +theoretical spectra and a spectrum identifier. We evaluate depths 4 to 10 at +a matching tolerance of 100 ppm. The result is the familiar list of +`SearchEngineResult` records, one per target and decoy spectrum, ranked by +descending score with the two delta fields filled in, as described on the +[SEQUEST-like page]({{root}}02_03_sequest_like_scoring.html). +*) + +let qMinAndMax = 4, 10 + +let andromedaResults = + AndromedaLike.calcAndromedaScore + qMinAndMax scanlimits 100. spectrum 20.93 2 precursorMZ theoSpecs "ms2Example" + +let printRanked (rs: SearchEngineResult.SearchEngineResult list) = + printfn "%-13s %-6s %8s %12s %8s" "sequence" "target" "score" "dBestToRest" "dNext" + rs + |> List.iter (fun r -> + printfn "%-13s %-6b %8.4f %12.4f %8.4f" + r.StringSequence r.IsTarget r.Score r.NormDeltaBestToRest r.NormDeltaNext) + +printRanked andromedaResults + +(** +```text +sequence target score dBestToRest dNext +ANLGMEVMHER true 86.4073 0.0000 0.2577 +AGGLGMEVMHER true 64.1383 0.2577 0.5796 +MANGLEVMHER true 14.0586 0.8373 0.0871 +EVMANLGMHER true 6.5320 0.9244 0.0756 +ANLGMEVMHER false 0.0000 1.0000 0.0000 +MANGLEVMHER false 0.0000 1.0000 0.0000 +EVMANLGMHER false 0.0000 1.0000 0.0000 +AGGLGMEVMHER false 0.0000 1.0000 0.0000 +``` + +The target of ANLGMEVMHER wins at 86.4, and the Gly-Gly variant is again +the runner-up ahead of both permutations, the same shape the SEQUEST-like +ranking had. The probability view separates the top pair more clearly, +though. Where the dot product saw a near-tie with a `dNext` of 0.013, the +binomial score puts a quarter of the top score between them. The mechanism +is visible in the model: a predicted peak that finds no partner enlarges n +without enlarging k and lowers the score, so the extra predictions of the +12 residue variant, and the few fragments it does not share with the true +ladder, cost it directly. All four decoys land at exactly zero: their raw +scores fall below the fixed correction terms explained next, and the clamp +cuts them off. + +## Reading an Andromeda-like score + +The score is derived from -10 log10 of the cumulative binomial probability. +On that scale a value of 60 corresponds to a probability of 10^-6 under the +random-match model. The implementation adds a mass-dependent correction +computed from the precursor m/z plus constant modification and cleavage +correction terms taken from the original Andromeda release, then subtracts +100. Negative results are clamped to zero. Absolute values are therefore +calibrated for ranking within this implementation. + +## Getting a hyperscore alongside + +`XScoring.calcAndromedaAndXTandemScore` runs the same rating and matching +machinery once per candidate and returns a pair of result lists, an +Andromeda-like ranking whose records carry `SearchEngine = AndromedaLike` +and an X!Tandem-like ranking carrying `SearchEngine = XTandemLike`, each +independently score-sorted with its own delta fields. It accepts the same +arguments and the same theoretical spectra as `calcAndromedaScore`. + +The X!Tandem score is the hyperscore: the summed intensity of all matched +measured peaks, taken as a logarithm, plus the log factorials of the number +of matched b ions and matched y ions, and a third log factorial for a separate +neutral-loss count. The factorials reward candidates that +match many ions of both series, since ten matched y ions weigh far more than +twice five. The counting is b/y-oriented, a matched peak contributes to the +count of the series flag it is tagged with, and since the loss peaks of this +library's generator carry their parent series flag, the separate neutral-loss +count stays empty here. The hyperscore is computed at +the deepest peak depth of the range, where the most measured peaks are +admitted to matching. +*) + +let andromedaResults2, hyperscoreResults = + XScoring.calcAndromedaAndXTandemScore + qMinAndMax scanlimits 100. spectrum 20.93 2 precursorMZ theoSpecs "ms2Example" + +printRanked hyperscoreResults + +(** +```text +sequence target score dBestToRest dNext +ANLGMEVMHER true 49.3979 0.0000 0.0994 +AGGLGMEVMHER true 44.4862 0.0994 0.5145 +MANGLEVMHER true 19.0709 0.6139 0.1745 +EVMANLGMHER true 10.4505 0.7884 0.1250 +EVMANLGMHER false 4.2767 0.9134 0.0082 +ANLGMEVMHER false 3.8712 0.9216 0.0000 +MANGLEVMHER false 3.8712 0.9216 0.0000 +AGGLGMEVMHER false 3.8712 0.9216 0.0000 +``` + +Both rankings agree on the order of the four targets. The decoys behave differently +under the hyperscore: a decoy still matches a few peaks by chance, and the logarithm +of their summed intensity enters the score even when the factorial terms contribute +nothing, so the decoys settle at a nonzero floor around 4 where the corrected +Andromeda score clamped them to zero. + +The Andromeda-like ranking returned alongside agrees with what +`AndromedaLike.calcAndromedaScore` produced on the same input, both modules +implement the same score. +*) + +printfn "%-13s %-6s %14s %9s" "sequence" "target" "AndromedaLike" "XScoring" +List.zip andromedaResults andromedaResults2 +|> List.iter (fun (a, x) -> + printfn "%-13s %-6b %14.4f %9.4f" a.StringSequence a.IsTarget a.Score x.Score) + +(** +```text +sequence target AndromedaLike XScoring +ANLGMEVMHER true 86.4073 86.4073 +AGGLGMEVMHER true 64.1383 64.1383 +MANGLEVMHER true 14.0586 14.0586 +EVMANLGMHER true 6.5320 6.5320 +ANLGMEVMHER false 0.0000 0.0000 +MANGLEVMHER false 0.0000 0.0000 +EVMANLGMHER false 0.0000 0.0000 +AGGLGMEVMHER false 0.0000 0.0000 +``` + +The score columns are identical for all eight records, so results from the +combined call can stand in for a separate `AndromedaLike` run. + +## Scoring whole runs + +For batch searches over many spectra the library also offers +`SearchEngineGeneric.OrderedCache.generateTheoSpectra`. It wires the +database lookup and the spectrum predictors together: given the ion series +calculator, the mass function, a peptide lookup function and three caches +(one for lookup results, one for Andromeda-style and one for SEQUEST-style +theoretical spectra), it answers a precursor mass window with the +theoretical spectra for both engine families, keeping previously generated +spectra in memory across consecutive windows and clearing the caches when a +memory ceiling is exceeded. The per-engine `getTheoSpecs` calls shown on +this page and the SEQUEST-like page are the direct route for scoring +individual spectra. + +## Where the scores go next + +The target and decoy scores and their deltas collected across a run feed +[false discovery rate control]({{root}}04_01_fdr_control.html), and the +identified peptides move on to +[quantification]({{root}}03_01_quantification.html). +*) From 1e28698fbe23b5de87bbff3f1e3ba00118a275e8 Mon Sep 17 00:00:00 2001 From: caroott Date: Mon, 31 Aug 2026 22:28:32 +0200 Subject: [PATCH 09/11] Add peak quantification documentation --- docs/03_01_quantification.fsx | 369 ++++++++++++++++++++++++++++++++++ 1 file changed, 369 insertions(+) create mode 100644 docs/03_01_quantification.fsx diff --git a/docs/03_01_quantification.fsx b/docs/03_01_quantification.fsx new file mode 100644 index 0000000..9217595 --- /dev/null +++ b/docs/03_01_quantification.fsx @@ -0,0 +1,369 @@ +(** +--- +title: Peak quantification +category: Quantification +categoryindex: 3 +index: 1 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Peak quantification + +The previous pages end with an identified peptide: the fragment spectrum recorded at +20.93 min matches ANLGMEVMHER, and its precursor sits at m/z 643.80 carrying two +charges (see [charge state determination]({{root}}01_03_charge_state_determination.html)). +Identification answers which peptide was measured. Quantification asks how much of it +was there, and the answer comes from a different slice through the data. + +A peptide elutes from the LC column over a stretch of retention time, together with +many other peptides that happen to leave the column at the same moment, and every +MS1 survey scan taken during that stretch records its isotope cluster once more. +All signals of one peptide ion species therefore form a three-dimensional feature +along m/z, retention time and intensity. Extracting the intensity at one m/z across +consecutive MS1 scans flattens that feature into the extracted ion chromatogram +(XIC): a trace that climbs while the peptide elutes and falls back to the noise +level once it has passed. The area +under that chromatographic peak is proportional to the amount of peptide, and area +integration is a more stable and reproducible quantity than the bare apex height. + +For the ANLGMEVMHER precursor one would extract the intensities around m/z 643.80 +from every MS1 scan of the run. The example data in this repository holds a single +MS1 scan, a single time point on such a trace. This page therefore builds a +realistic XIC trace synthetically, which has an advantage for a walkthrough: the +generating parameters are known exactly, so every result can be checked against +ground truth. + +## Building an XIC trace + +XICs approximately follow a Gaussian shape, with one recurring deviation: peak +tailing, a positive skew commonly caused by column overloading or unwanted +retention mechanisms that keep part of the analyte on the column a little longer. +The exponentially modified Gaussian (EMG) describes such tailing peaks well, so the +main synthetic peak is generated from exactly that function. A second, smaller feature elutes 0.85 min later, and +seeded uniform noise imitates the positive chemical background an instrument +records between peaks. +*) + +open System +open FSharp.Stats +open BioFSharp.Mz +open BioFSharp.Mz.Quantification + +// Exponentially modified Gaussian: a Gaussian of height amplitude, center meanX +// and width sigma, convolved with an exponential decay of relaxation time tau. +// The tau term produces the right-sided tail typical of chromatographic peaks. +let emg amplitude meanX sigma tau x = + (amplitude * sigma / tau) * sqrt (Math.PI / 2.) + * exp (0.5 * (sigma / tau) ** 2. - (x - meanX) / tau) + * SpecialFunctions.Errorfunction.Erfc ((sigma / tau - (x - meanX) / sigma) / sqrt 2.) + +let gaussian amplitude meanX sigma x = + amplitude * exp (-((x - meanX) ** 2.) / (2. * sigma ** 2.)) + +// Fixed seed: reruns of this page produce identical numbers. +let rnd = Random(1337) + +// One MS1 scan every 0.7 s (0.0117 min) for three minutes around the 20.9 min +// elution of the running example. The main peak reaches the tens of thousands +// of counts, a realistic MS1 XIC intensity for a well ionizing peptide, and its +// sigma of 0.055 min (3.3 s) is a typical chromatographic peak width. The +// noise floor of up to 250 counts stays strictly positive, as measured +// intensities do. +let retentionTimes = [| 19.6 .. 0.0117 .. 22.6 |] + +let intensities = + retentionTimes + |> Array.map (fun rt -> + emg 42000. 20.9 0.055 0.09 rt + + gaussian 9000. 21.75 0.05 rt + + rnd.NextDouble() * 250.) + +let apexIndex = + intensities |> Array.findIndex ((=) (Array.max intensities)) + +printfn "%i points from %.2f to %.2f min" retentionTimes.Length retentionTimes.[0] (Array.last retentionTimes) +printfn "highest intensity: %.0f at %.3f min" intensities.[apexIndex] retentionTimes.[apexIndex] + +(** +```text +257 points from 19.60 to 22.60 min +highest intensity: 27510 at 20.946 min +``` + +The apex lands a little right of the EMG center at 20.9 min because the exponential +tail shifts the maximum, exactly as it does on a real column. The generating EMG has +an analytic area of amplitude times sigma times the square root of two pi, about +5790 intensity·min, a number to keep in mind for the results below. + +## Detecting peaks on the trace + +Quantification starts by deciding where on the trace a peak begins and ends. +FSharp.Stats brings a second-derivative peak picker for this, +`Signal.PeakDetection.SecondDerivative.getPeaks`. It smooths the trace with a +Savitzky-Golay filter, takes the negative second derivative (which turns every +peak into a pronounced maximum), estimates the noise as the mean absolute +difference between the raw and the smoothed trace, and keeps candidate peaks that +rise above a signal-to-noise cutoff times that estimate. Its arguments are the +cutoff followed by the polynomial order and the window width in points of the +Savitzky-Golay filter. +*) + +let peaks = + Signal.PeakDetection.SecondDerivative.getPeaks 5. 2 13 retentionTimes intensities + +peaks +|> Array.iter (fun p -> + printfn "apex %.3f min, intensity %.0f, from %.3f to %.3f min, %i points" + p.Apex.XVal p.Apex.YVal p.LeftEnd.XVal p.RightEnd.XVal p.XData.Length) + +(** +```text +apex 20.946 min, intensity 27510, from 20.712 to 21.495 min, 68 points +apex 21.753 min, intensity 9114, from 21.542 to 21.917 min, 33 points +``` + +Both synthetic features are found and nothing else is. The cutoff carries that +result: with 0.5 instead of 5 the same call reports 15 peaks, most of them noise +ripples of around 200 counts. Each detected peak is an `IdentifiedPeak` holding the +apex, the left and right peak ends, the lift-off points where curvature indicates +the peak rising out of the baseline, and the slice of the measured data between the +ends. Note how the first peak's slice reaches from 20.71 to 21.50 min, well beyond +a symmetric window around the apex, because the border search followed the tail. +This record is the input type the quantification below consumes. + +## Detecting peaks with the chromatographic wavelet + +This library also ships its own chromatographic peak detector, +`PeakDetection.Wavelet.identifyPeaks`, which correlates the trace with Mexican Hat +(Ricker) wavelets of increasing scale. The Mexican Hat is the negative normalized +second derivative of a Gaussian, so it matches the shape a chromatographic peak is +expected to have, and shape matching lets weak peaks stand out against noise of +similar amplitude. We hand the detector the trace on a seconds axis, where the +0.7 s sampling and peak widths of a few seconds give the wavelet a convenient +scale range. Two parameters deserve a note. `Borderpadding` must be generous +relative to the wavelet width, and 200 works well on traces of this size. +`MaxPeakLength` caps the widest wavelet scale at one sixth of its value, and 30 s +sits comfortably above six times the 3.3 s sigma of the synthetic peak. +*) + +let traceInSeconds = + Array.map2 (fun rt intensity -> rt * 60., intensity) retentionTimes intensities + +let waveletParameters : PeakDetection.Wavelet.Parameters = + { Borderpadding = 200 + BorderPadMethod = Signal.Padding.BorderPaddingMethod.Zero + InternalPaddingMethod = Signal.Padding.InternalPaddingMethod.LinearInterpolation + HugeGapPaddingMethod = Signal.Padding.HugeGapPaddingMethod.Zero + HugeGapPaddingDistance = 100. + MaxPeakLength = 30. + NoiseQuantile = 0.5 + MinSNR = 1. } + +let peakGroups = + PeakDetection.Wavelet.identifyPeaks waveletParameters traceInSeconds + +let coveredByGroup rtInMinutes = + peakGroups + |> List.exists (fun g -> g.Start <= rtInMinutes * 60. && rtInMinutes * 60. <= g.End) + +printfn "peak groups found: %i" peakGroups.Length +printfn "a group covers the main apex: %b" (coveredByGroup peaks.[0].Apex.XVal) +printfn "a group covers the second apex: %b" (coveredByGroup peaks.[1].Apex.XVal) + +(** +```text +peak groups found: 29 +a group covers the main apex: true +a group covers the second apex: true +``` + +The detector finds both true apexes, and it detects generously: on this two-feature +trace it reports 29 peak groups, treating shoulders and noise structures next to the +real peaks as separate groups. A consumer therefore selects the group covering the +retention time of interest and disregards the rest. A group's `Data` holds the +padded trace region the wavelet worked on, and the fitted `Stdev` reflects the +wavelet scale the group responded to within the configured range. For the rest of +this page we stay with the second-derivative peaks, which carry exact slice +boundaries and feed directly into the quantification step. + +## Fitting and integrating the main peak + +The `HULQ` module turns an `IdentifiedPeak` into a quantity. `HULQ.getPeakBy` +selects from the detected peaks the one whose boundaries contain a target retention +time, falling back to the nearest apex when no peak contains it. The natural target +is the retention time of the identification, 20.93 min for our MS2 scan. + +`HULQ.quantifyPeak` then runs the model fitting workflow. It estimates starting +parameters from the peak slice (by weighted moments, with a Caruana log-parabola +fit stepping in when the slice is truncated on one side), fits a Gaussian and, when +the moments allow, an EMG via Levenberg-Marquardt, scores both fits by their +standard error of prediction, keeps the better one, and integrates the fitted model +analytically for the area. +*) + +let describeModel (q: HULQ.QuantifiedPeak) = + match q.Model with + | Some (HULQ.Gaussian _) -> "Gaussian" + | Some (HULQ.EMG _) -> "EMG" + | None -> "none, trapezoid fallback" + +let printQuantification (q: HULQ.QuantifiedPeak) = + printfn "model selected : %s" (describeModel q) + match q.Model, q.EstimatedParams with + | Some (HULQ.Gaussian _), [| amp; meanX; sigma |] -> + printfn "amplitude : %.1f" amp + printfn "mean retention time : %.3f min" meanX + printfn "sigma : %.4f min" sigma + | Some (HULQ.EMG _), [| amp; meanX; sigma; tau |] -> + printfn "amplitude : %.1f" amp + printfn "mean retention time : %.3f min" meanX + printfn "sigma : %.4f min" sigma + printfn "tau : %.4f min" tau + | _ -> () + printfn "standard error of prediction: %.1f" q.StandardErrorOfPrediction + printfn "area under the fitted model : %.1f" q.Area + printfn "measured apex intensity : %.1f" q.MeasuredApexIntensity + +let mainPeak = HULQ.getPeakBy peaks 20.93 + +let mainQuantification = HULQ.quantifyPeak mainPeak + +printQuantification mainQuantification + +(** +```text +model selected : EMG +amplitude : 35762.5 +mean retention time : 20.906 min +sigma : 0.0629 min +tau : 0.0759 min +standard error of prediction: 601.2 +area under the fitted model : 5635.9 +measured apex intensity : 27509.6 +``` + +The tailed peak is recognized as an EMG. The fitted mean retention time of +20.906 min sits on the generating center of 20.9, and sigma and tau land near the +generating 0.055 and 0.09 min. The amplitude is the height parameter of the +pre-convolution Gaussian, so it neither matches the measured apex of 27510 counts +nor needs to. The `MeasuredApexIntensity` field preserves that raw apex alongside +the model. + +How good is the model area? A direct numerical integration of the same slice gives +an assumption-free reference. +*) + +let mainTrapezoid = + Integration.trapezEstAreaOf mainPeak.XData mainPeak.YData + +printfn "model area : %.1f" mainQuantification.Area +printfn "trapezoid area: %.1f" mainTrapezoid + +(** +```text +model area : 5635.9 +trapezoid area: 5874.7 +``` + +Both land close to the analytic area of 5790 intensity·min the trace was generated +with. The trapezoid integral runs a little above it because it also sums the noise +floor riding on the slice, the model area a little below. The model area is the +value a pipeline stores: it is anchored to a fitted peak shape, which keeps it +comparable when slice boundaries or noise differ between runs. + +## How the model is chosen + +Whether an EMG is attempted at all is decided by the moment estimates. +`ParameterEstimation.estTau` derives the EMG tail parameter from the moments as +sigma times the cube root of half the skewness. For a peak with zero or negative +skew that fractional power of a non-positive number is NaN, the EMG candidate +becomes unavailable, and the Gaussian is used. That is the intended selection path: +a peak that is not right-skewed is a Gaussian case, and no tail parameter should be +invented for it. The moments of our two peaks show both sides of this rule. +*) + +let secondPeak = HULQ.getPeakBy peaks 21.75 + +let printMoments name peak = + match ParameterEstimation.estimateMoments peak with + | Some m -> + printfn "%s: mean %.3f min, sigma %.4f min, skew %.3f, tau %.4f" name m.MeanX m.Std m.Skew m.Tau + | None -> printfn "%s: no moment estimate" name + +printMoments "main peak " mainPeak +printMoments "second peak" secondPeak + +(** +```text +main peak : mean 20.991 min, sigma 0.1068 min, skew 1.109, tau 0.0878 +second peak: mean 21.748 min, sigma 0.0555 min, skew -0.212, tau NaN +``` + +The tailed main peak carries a clear positive skew and a defined tau, so both +models compete and the output above showed the EMG winning on standard error. The +second peak was generated symmetric, its slight negative skew is noise, and its tau +is NaN. For it the EMG leg never runs. + +## Quantifying the second feature + +The workflow repeats per peptide feature: select the peak at the feature's +retention time, quantify it. +*) + +let secondQuantification = HULQ.quantifyPeak secondPeak + +printQuantification secondQuantification + +(** +```text +model selected : Gaussian +amplitude : 9032.7 +mean retention time : 21.749 min +sigma : 0.0516 min +standard error of prediction: 121.0 +area under the fitted model : 1167.3 +measured apex intensity : 9113.5 +``` + +As the moments predicted, the symmetric feature is fitted as a Gaussian, and for a +Gaussian the amplitude is the apex height, so 9033 sits right at the generating +9000. Mean and sigma recover the generating 21.75 min and 0.05 min. Should neither +model converge on some degenerate slice, `quantifyPeak` still returns a usable +result, the trapezoid area of the slice with `Model = None`. + +## From one peak to a pipeline + +The error measure behind the model selection is +`NonLinearRegression'.standardErrorOfPrediction` from this library's FSharp.Stats +extension, the root of the summed squared residuals normalized by the residual +degrees of freedom, and the model with the lower value wins. Around this core a +real pipeline adds the comparative layer: for isotopic labeling such as 15N, the +light and the heavy XIC of the same peptide are extracted and their fitted areas +are ratioed within one run, while label-free workflows compare the areas of +matching features across runs. + +Before peak areas are aggregated into protein quantities, the identifications they +hang on are filtered to a controlled error rate, which is the subject of +[FDR control]({{root}}04_01_fdr_control.html). And the m/z every XIC follows comes +from the precursor's charge assignment on the +[charge state determination]({{root}}01_03_charge_state_determination.html) page. +*) From 6577a6311f1da5e0f71663fa68542c91a24e0f86 Mon Sep 17 00:00:00 2001 From: caroott Date: Tue, 1 Sep 2026 18:55:43 +0200 Subject: [PATCH 10/11] Add false discovery rate control documentation --- docs/04_01_fdr_control.fsx | 346 +++++++++++++++++++++++++++++++++++++ 1 file changed, 346 insertions(+) create mode 100644 docs/04_01_fdr_control.fsx diff --git a/docs/04_01_fdr_control.fsx b/docs/04_01_fdr_control.fsx new file mode 100644 index 0000000..3491efd --- /dev/null +++ b/docs/04_01_fdr_control.fsx @@ -0,0 +1,346 @@ +(** +--- +title: False discovery rate control +category: Validation and inference +categoryindex: 4 +index: 1 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# False discovery rate control + +Over a whole run, the per-spectrum rankings of the scoring pages +([SEQUEST-like]({{root}}02_03_sequest_like_scoring.html) and +[Andromeda-like]({{root}}02_04_andromeda_like_scoring.html)) accumulate into a long +list of peptide spectrum matches (PSMs), each carrying a score. +The score states how well a spectrum and a candidate agree. It says nothing +about how often agreement this good arises by chance, so a score alone +cannot separate true identifications from false ones. The search database +never covers every peptide actually present in the sample, so for some +spectra the true answer was not among the candidates and the best available +match is wrong. The instrument also sometimes selects precursor ions that +were never peptides. And in a large search space, some wrong candidate will +score well on some spectrum simply because so many candidates were tried. + +The standard way to put numbers on this problem is the decoy database. +Reversed or shuffled sequences are searched alongside the real ones, and any +PSM that lands on a decoy is wrong by construction. Counting decoy hits +above a score threshold therefore estimates how many target hits above the +same threshold are wrong, because a wrong target assignment behaves like a +decoy assignment. + +Three quantities are built on that count. The global false discovery rate +(FDR) of an accepted PSM list is the fraction of the list that is wrong. The +q-value attached to a score is the FDR of the list you get by accepting +everything at that score or better, made monotone so that relaxing the +threshold never claims a lower error. It is the number used for cutoffs, and +"filter at 1 percent FDR" means keeping PSMs with q at or below 0.01. The +posterior error probability (PEP) asks the question for a single PSM: how +likely is this one match wrong. Tools in this space include Qvality, which +estimates PEPs and q-values by regression on decoy counts, and Percolator, +which rescores PSMs with an iteratively trained SVM and appears again at the +end of this page. The `FDRControl` module implements the counting and +fitting machinery for q-values. + +## Simulating a run's worth of PSM scores + +The scoring pages worked on one spectrum, so for this page we simulate the +score list of a whole run. A seeded generator and a Box-Muller helper draw +normally distributed scores on an Andromeda-like scale from 0 to about 120. +*) + +open BioFSharp.Mz + +let rng = System.Random 42 + +// Box-Muller transform: two uniform draws become one normal draw. +let gaussian mean sd = + let u1 = 1.0 - rng.NextDouble() + let u2 = rng.NextDouble() + mean + sd * (sqrt (-2.0 * log u1) * cos (2.0 * System.Math.PI * u2)) + +// Decoy PSMs are wrong by construction and score low. +let decoyScores = Array.init 5000 (fun _ -> max 0.0 (gaussian 25.0 8.0)) + +// Target PSMs are a mixture. 60 percent are correct identifications and +// score high, 40 percent are wrong assignments. The wrong targets draw from +// the same low distribution as the decoys, because that equivalence is the +// premise of the method: the decoy score distribution estimates how wrong +// target assignments score. +let correctScores = Array.init 3000 (fun _ -> max 0.0 (gaussian 70.0 15.0)) +let wrongScores = Array.init 2000 (fun _ -> max 0.0 (gaussian 25.0 8.0)) +let targetScores = Array.append correctScores wrongScores + +let psms = + Array.append + (targetScores |> Array.map (fun s -> FDRControl.createQValueInput s false)) + (decoyScores |> Array.map (fun s -> FDRControl.createQValueInput s true)) + +let median xs = + let sorted = Array.sort xs + sorted.[Array.length sorted / 2] + +printfn "PSMs: %i targets, %i decoys" targetScores.Length decoyScores.Length +printfn "median target score: %.1f median decoy score: %.1f" + (median targetScores) (median decoyScores) + +(** +```text +PSMs: 5000 targets, 5000 decoys +median target score: 56.4 median decoy score: 25.1 +``` + +`createQValueInput` builds the `QValueInput` record the module works with, a +score plus an `IsDecoy` flag. The functions below take the data together +with three accessors, one for the decoy flag and one score projection each +for decoys and targets. Here both projections read the same field. + +## Computing q-values by counting decoys + +`calculateQValueStorey` implements the direct counting estimate and returns +a function from score to q-value. Internally it walks the scores from best +to worst and accumulates decoy and target counts. At each score the raw +estimate is accumulated decoys divided by accumulated targets. A second pass +makes the sequence monotone, so q never decreases as the score drops, and a +linear interpolation over the resulting support turns the table into a +function usable at any score. +*) + +let storeyQ = + FDRControl.calculateQValueStorey + psms + (fun x -> x.IsDecoy) + (fun x -> x.Score) + (fun x -> x.Score) + +let probes = [90.; 70.; 55.; 40.] + +for s in probes do + printfn "score %5.0f q = %.5f" s (storeyQ s) + +(** +```text +score 90 q = 0.00000 +score 70 q = 0.00000 +score 55 q = 0.00000 +score 40 q = 0.05299 +``` + +No decoy in this simulated run scores above 55, so down to there the +accumulated decoy count is zero and so is q. By 40 the upper tail of the +decoy distribution is contributing steadily and q has passed 5 percent. +Accepting at a q threshold is a score cutoff in disguise, so counting the +targets that pass is one filter away. +*) + +let acceptedTargets threshold = + targetScores |> Array.filter (fun s -> storeyQ s <= threshold) |> Array.length + +printfn "targets accepted at q <= 0.01: %i" (acceptedTargets 0.01) +printfn "targets accepted at q <= 0.05: %i" (acceptedTargets 0.05) +printfn "wrong targets among the q <= 0.01 list: %i" + (wrongScores |> Array.filter (fun s -> storeyQ s <= 0.01) |> Array.length) + +(** +```text +targets accepted at q <= 0.01: 2897 +targets accepted at q <= 0.05: 3003 +wrong targets among the q <= 0.01 list: 6 +``` + +Against ground truth, the 1 percent list holds 2897 PSMs of which 6 are actually +wrong, a true error rate of 0.2 percent. The estimate is conservative here because all 5000 +decoys stand in for the 2000 wrong targets, so the decoy count overstates +the false hits. The assumed fraction of wrong targets is exactly what the +pi0 parameter of the next method makes explicit. + +## Smoothing the estimate with a logistic fit + +The counting estimate is a step function shaped by every random fluctuation +in the decoy tail. `calculateQValueLogReg` trades the steps for a smooth +curve. First `binningFunction` slices the score axis into bandwidth-wide +bins and computes raw estimates per bin. The decoy count is normalized by +the overall decoy share, then doubled and scaled by pi0. Dividing that by the +total count in the bin gives a PEP-like value, and dividing it by the total +count at this bin and above gives a q-like value. These raw bin values serve +purely as input points for the fit. A descending logistic curve is then fitted through the +(score, q) points with constrained Levenberg-Marquardt over a range of +steepness guesses, and the fitted curve is the deliverable, a smooth +function from score to q. + +The first argument is pi0, the assumed fraction of wrong assignments among +the targets. We pass 0.4 because the generated mixture is 40 percent wrong, +which is what a real analysis would estimate from the score overlap. The +bandwidth of 3 gives bins a few score units wide, enough entries per bin on +a 0 to 120 scale to keep the raw points stable. +*) + +let logRegQ = + FDRControl.calculateQValueLogReg + 0.4 + 3.0 + psms + (fun x -> x.IsDecoy) + (fun x -> x.Score) + (fun x -> x.Score) + +printfn "%5s %10s %10s" "score" "Storey q" "logReg q" +for s in probes do + printfn "%5.0f %10.5f %10.5f" s (storeyQ s) (logRegQ s) + +(** +```text +score Storey q logReg q + 90 0.00000 0.00000 + 70 0.00000 0.00000 + 55 0.00000 0.00000 + 40 0.05299 0.00065 +``` + +The methods agree over most of the range. Everywhere above 55 both report +zero, and both place the onset of error below 45. They part ways in the +transition zone: at a score of 40 the counting estimate already sees the +159 decoys above that score, while the fitted sigmoid centers its drop on +the bulk of the decoy distribution near 30 and has fallen close to zero by +40. The pi0 scaling also lowers the fitted curve as a whole, since the fit +treats only 40 percent of the targets as potentially wrong where Storey's +count treats every decoy as evidence against a target. Which output feeds +the pipeline is a choice between the assumption-free step function and the +smooth curve that needs a pi0. + +## From list-level q to single-PSM PEP + +The q-value describes a whole accepted list. The PEP asks how likely one particular +PSM is wrong. A PSM +sitting exactly at a 1 percent q cutoff typically carries a PEP well above 1 +percent, because the list average is pulled down by all the confident hits +above it. The library derives PEP-like values inside the binning and fitting +machinery you just saw, where each bin's decoy share is a local error +estimate, and it also exposes dedicated PEP functions: `calculatePEPValues` +computes the per-bin decoy shares directly and `initCalculateLin` returns a +fitted score-to-PEP mapping. The pipeline-facing outputs on this page are the +two q-value functions. + +## Estimating protein-level false positives with MAYU + +Once PSMs are aggregated to proteins, the error question returns one level +up: how many of the target proteins with at least one accepted hit are +false? `FDRControl.MAYU.estimatePi0HG` answers with the MAYU model, +translated from Percolator's ProteinFDREstimator. It treats the decoy +protein hits as draws from a hypergeometric distribution over the database +and returns the expected number of false positives among the target hits. +The arguments are the total number of candidate proteins in the database, +the number of target protein hits and the number of decoy protein hits. +*) + +let expectedFP = FDRControl.MAYU.estimatePi0HG 2000. 800. 30. + +printfn "expected false positive protein hits: %.2f" expectedFP +printfn "implied protein-level FDR: %.4f" (expectedFP / 800.) + +(** +```text +expected false positive protein hits: 18.27 +implied protein-level FDR: 0.0228 +``` + +With 800 of 2000 database proteins hit and 30 decoy hits observed, the model +expects about 18 of the 800 target hits to be false, a protein-level FDR +just over 2 percent. The expectation is a weighted average over all feasible +false positive counts from zero up to the decoy count, so it can land below +the raw decoy count, as it does here. The [protein inference]({{root}}04_02_protein_inference.html) +page applies this estimator when assembling peptide evidence into protein +lists. + +## Rescoring with Percolator + +The methods above interpret one score column. Percolator improves the +separation before the interpretation: it trains a support vector machine on +the PSMs themselves, using the decoy PSMs as the negative training set and +high-confidence target PSMs as positives, on features such as the raw +score, the delta to the next-best candidate, the precursor mass error and +the precursor charge. The learned score replaces the raw one and the +positive training set is re-derived from it, and this classification round +repeats until the score stabilizes. The output is again q-values and PEPs, +computed on a score axis where targets and decoys separate better. + +Percolator ships as an external program, and the `PercolatorWrapper` module +wraps its command line. The option groups of the CLI are mirrored by +discriminated unions under `PercolatorWrapper.Parameters`, so a parameter +set is ordinary typed F# data, and the `PercolatorParams` cases collect the +groups. Building a set looks like this. +*) + +open BioFSharp.Mz.PercolatorWrapper + +let percolatorParams : Parameters.PercolatorParams list = + [ Parameters.GeneralOptions + [ Parameters.VerbosityOfOutput 2 + Parameters.PostProcessing_TargetDecoyCompetition ] + Parameters.FileInputOptions + [ Parameters.PINTAB (System.IO.FileInfo "run1.pin") ] + Parameters.FileOutputOptions + [ Parameters.POUTTAB_PSMs (System.IO.FileInfo "run1_psms.tsv") + Parameters.POUTTAB_DecoyPSMs (System.IO.FileInfo "run1_decoy_psms.tsv") ] + Parameters.SVMTrainingOptions + [ Parameters.FDR_CrossValidation 0.01 + Parameters.MaxIterations 10 ] ] + +let groupSummary (p: Parameters.PercolatorParams) = + match p with + | Parameters.GeneralOptions s -> "general options", Seq.length s + | Parameters.FileInputOptions s -> "file input", Seq.length s + | Parameters.FileOutputOptions s -> "file output", Seq.length s + | Parameters.SVMFeatureOptions s -> "SVM features", Seq.length s + | Parameters.SVMTrainingOptions s -> "SVM training", Seq.length s + | Parameters.ProteinInferenceOptions_FIDO s -> "Fido inference", Seq.length s + | Parameters.ProteinInferenceOptions_Percolator s -> "protein inference", Seq.length s + +for p in percolatorParams do + let name, count = groupSummary p + printfn "%-16s %i option(s)" name count + +(** +```text +general options 2 option(s) +file input 1 option(s) +file output 2 option(s) +SVM training 2 option(s) +``` + +The set reads a pin-tab input file, applies target-decoy competition, +evaluates cross validation at 1 percent FDR with at most ten SVM iterations, +and writes separate tab-delimited result files for target and decoy PSMs. +Running the set goes through `PercolatorWrapper(os).Percolate`, which +renders the options to a command line for the chosen operating system and +launches the percolator process. Executing it requires an external +percolator installation, so this page stops at the constructed parameters. + +## Where the identifications go next + +Whichever q-value function the pipeline uses, the PSMs that pass the chosen +threshold form the statistically controlled identification list. Those +identifications carry their peptide evidence into +[protein inference]({{root}}04_02_protein_inference.html), and the ion areas +determined by [quantification]({{root}}03_01_quantification.html) are +aggregated over the same accepted set. +*) From 67f8219f42fd4adb18dd97eed51a5c4c36275c3e Mon Sep 17 00:00:00 2001 From: caroott Date: Tue, 1 Sep 2026 19:16:33 +0200 Subject: [PATCH 11/11] Add protein inference documentation --- docs/04_02_protein_inference.fsx | 532 +++++++++++++++++++++++++++++++ 1 file changed, 532 insertions(+) create mode 100644 docs/04_02_protein_inference.fsx diff --git a/docs/04_02_protein_inference.fsx b/docs/04_02_protein_inference.fsx new file mode 100644 index 0000000..5cea664 --- /dev/null +++ b/docs/04_02_protein_inference.fsx @@ -0,0 +1,532 @@ +(** +--- +title: Protein inference +category: Validation and inference +categoryindex: 4 +index: 2 +--- +*) + +(*** hide ***) + +(*** condition: prepare ***) +#r "nuget: FSharpAux, 2.1.0" +#r "nuget: FSharpAux.IO, 2.1.0" +#r "nuget: FSharp.Stats, 0.6.0" +#r "nuget: BioFSharp, 2.0.0" +#r "nuget: Plotly.NET, 6.0.0-preview.2" +#r "../src/BioFSharp.Mz/bin/Release/netstandard2.0/BioFSharp.Mz.dll" +#r "../src/BioFSharp.Mz.Vis/bin/Release/netstandard2.0/BioFSharp.Mz.Vis.dll" + +(*** condition: ipynb ***) +#if IPYNB +#r "nuget: BioFSharp.Mz, {{fsdocs-package-version}}" +#r "nuget: BioFSharp.Mz.Vis, {{fsdocs-package-version}}" +#endif // IPYNB + +(** +[![Binder]({{root}}img/badge-binder.svg)](https://mybinder.org/v2/gh/BioFSharp/BioFSharp.Mz/gh-pages?filepath={{fsdocs-source-basename}}.ipynb)  +[![Script]({{root}}img/badge-script.svg)]({{root}}{{fsdocs-source-basename}}.fsx)  +[![Notebook]({{root}}img/badge-notebook.svg)]({{root}}{{fsdocs-source-basename}}.ipynb) + +# Protein inference + +The [FDR control]({{root}}04_01_fdr_control.html) page ended with a +statistically controlled list of accepted peptide identifications. +Biological questions are asked about proteins, and mapping the accepted +peptides back onto proteins runs into an ambiguity. A tryptic peptide often +occurs in more than one protein of the database. Splice variants of a gene +share most of their coding sequence, and gene families carry conserved +stretches that digest into identical peptides. When such a peptide is +observed, the evidence supports every protein that contains it, and nothing +in the measurement can tell those proteins apart. Protein inference resolves +this by reporting protein groups: sets of accessions that the observed +peptides cannot distinguish, carried through the rest of the pipeline as one +entry. + +The `ProteinInference` module describes each observed peptide as a +`ProteinClassItem`, which couples the peptide sequence with the array of +protein accessions it maps to and a `PeptideEvidenceClass`. The class states +how specifically the peptide points at a gene model. `C1a` marks a peptide +that maps to exactly one splice variant of one gene. `C1b` marks one that +maps to several splice variants which translate to the same protein +sequence. `C2a` and `C2b` cover peptides shared between splice variants that +differ elsewhere, either a subset of them or all of them. `C3a` and `C3b` +cover peptides shared between different genes, where `C3a` is the case of +genes encoding an identical protein sequence and `C3b` the general case. +Inference processes the classes in that order, and in the output every group +carries the class of the evidence that created it. + +In a full pipeline these classes come from genome annotation. The GFF3 +helpers of the module read an annotation file: `assignTranscriptsToGenes` +collects the mRNA entries under their gene locus, with +`createProteinModelInfoFromEntry` extracting the splice variant identifier +of each transcript, and `createPeptideProteinRelation` maps the peptides of +an in silico digest onto the resulting gene models. From that relation the +evidence class of every peptide follows from how it distributes over loci +and splice variants. This page skips the annotation step and constructs the +class items directly, which keeps the example self-contained. + +## Describing the peptide evidence + +The example imitates a small slice of a Chlamydomonas reinhardtii +measurement, built so that every interesting mapping case occurs at least +once. Two chloroplast proteins have only unique peptides. A light-harvesting +gene has two annotated splice variants and every observed peptide sits in +their shared exons, so those peptides map to both variants. Two actin-like +paralogs each have a unique peptide and additionally share one conserved +peptide. One protein is a one-hit wonder with a single weakly scoring +peptide, which becomes important once decoy scores enter. +*) + +open BioFSharp.Mz +open BioFSharp.PeptideClassification + +let items = + [ + // psbA (photosystem II D1) and rbcL (rubisco large subunit): two + // unique peptides each, mapping to exactly one gene model (C1a). + ProteinInference.createProteinClassItem [|"psbA"|] PeptideEvidenceClass.C1a "VLNTWADIINR" + ProteinInference.createProteinClassItem [|"psbA"|] PeptideEvidenceClass.C1a "ETTENESANEGYR" + ProteinInference.createProteinClassItem [|"rbcL"|] PeptideEvidenceClass.C1a "DTDILAAFR" + ProteinInference.createProteinClassItem [|"rbcL"|] PeptideEvidenceClass.C1a "LTYYTPDYVVR" + // A light-harvesting gene with two splice variants. Both observed + // peptides lie in shared exons and map to all isoforms of the gene + // (C2b), so the two variants are indistinguishable here. + ProteinInference.createProteinClassItem + [|"Cre01.g001000.t1"; "Cre01.g001000.t2"|] PeptideEvidenceClass.C2b "SSYLDGSAPGDFGFDPLGLGK" + ProteinInference.createProteinClassItem + [|"Cre01.g001000.t1"; "Cre01.g001000.t2"|] PeptideEvidenceClass.C2b "NLAGDVIGTR" + // Two actin-like paralogs: each has one unique peptide (C1a) ... + ProteinInference.createProteinClassItem [|"Cre02.g095150.t1"|] PeptideEvidenceClass.C1a "SYELPDGQVITIGNER" + ProteinInference.createProteinClassItem [|"Cre03.g144807.t1"|] PeptideEvidenceClass.C1a "VAPEEHPVLLTEAPLNPK" + // ... and they share one conserved peptide across the two gene loci + // (C3b, different genes with different protein sequences). + ProteinInference.createProteinClassItem + [|"Cre02.g095150.t1"; "Cre03.g144807.t1"|] PeptideEvidenceClass.C3b "AGFAGDDAPR" + // A one-hit wonder: a single peptide, and its PSM score will be low. + ProteinInference.createProteinClassItem [|"Cre06.g250100.t1"|] PeptideEvidenceClass.C1a "LISWYDNEWGYSNR" + ] + +let distinctProteins = + items + |> List.collect (fun item -> List.ofArray item.GroupOfProteinIDs) + |> List.distinct + +printfn "%i peptides mapping onto %i proteins" items.Length distinctProteins.Length + +(** +```text +10 peptides mapping onto 7 proteins +``` + +## Grouping the evidence into protein groups + +`inferSequences` turns the class items into `InferredProteinClassItem` values, one +per protein group, each recording the group's evidence class and the peptides kept +for its quantification. The `IntegrationStrictness` decides what happens to +overlapping groups: `Minimal` merges them down to the smallest set of proteins that +still explains all observed peptides, while `Maximal` keeps every evidence group +exactly as the peptides stated it. The `PeptideUsageForQuantification` decides which +peptides a finished group keeps: under `Minimal` only those, among all peptides +consistent with it, whose own protein set is smallest and therefore points at the +group most specifically. Under `Maximal` it keeps every peptide whose protein set +contains the group, and under `MaximalInverse` every peptide whose protein set is +contained in it. The production settings of this module are `Maximal` strictness +with `Minimal` peptide usage, and the call writes a progress line to standard output +while it assembles its lookup structures. +*) + +let inferred = + ProteinInference.inferSequences + ProteinInference.IntegrationStrictness.Maximal + ProteinInference.PeptideUsageForQuantification.Minimal + items + |> List.ofSeq + +(** +```text +finish setup +``` + +That progress marker is everything the call itself prints under `Maximal` +strictness. The result prints as one line per protein group. +`proteinGroupToString` joins the accessions of a group with semicolons, and +`inferSequences` already returns that joined representation. +*) + +for g in inferred do + printfn "%-33s %s %s" + g.GroupOfProteinIDs + (sprintf "%A" g.Class) + (String.concat ", " g.PeptideSequence) + +(** +```text +psbA C1a VLNTWADIINR, ETTENESANEGYR +rbcL C1a DTDILAAFR, LTYYTPDYVVR +Cre02.g095150.t1 C1a SYELPDGQVITIGNER +Cre03.g144807.t1 C1a VAPEEHPVLLTEAPLNPK +Cre06.g250100.t1 C1a LISWYDNEWGYSNR +Cre01.g001000.t1;Cre01.g001000.t2 C2b SSYLDGSAPGDFGFDPLGLGK, NLAGDVIGTR +Cre02.g095150.t1;Cre03.g144807.t1 C3b AGFAGDDAPR +``` + +The two splice variants collapsed into a single group, because no observed +peptide separates them. The actin-like paralogs stayed apart, since each has +unique evidence, and their shared peptide formed a composite group of its +own. `Maximal` strictness meant that every evidence group was reported +unchanged, seven groups in total. The `Minimal` peptide usage shows in the +paralog groups: the shared peptide `AGFAGDDAPR` was consistent with the +`Cre02` group as well, but only the unique peptide points at that group +most specifically, so only the unique peptide is kept for quantifying it. + +For comparison, the same evidence under `Minimal` strictness. On this path +the merge logic runs and traces its decisions to standard output, so the +call prints a diagnostic line for every piece of evidence that overlaps +nothing recorded so far and echoes the existing group when further evidence +arrives on the same proteins. +*) + +let inferredMinimal = + ProteinInference.inferSequences + ProteinInference.IntegrationStrictness.Minimal + ProteinInference.PeptideUsageForQuantification.Minimal + items + |> List.ofSeq + +(** +```text +Search did not return anything +psbA +Search did not return anything +rbcL +Search did not return anything +Search did not return anything +Search did not return anything +Search did not return anything +Cre01.g001000.t1;Cre01.g001000.t2 +finish setup +``` + +Every group is announced once when its first evidence arrives, and the +echoed group names accompany the integration of the second peptide on the +same proteins. The last line is the same progress marker as above, and the +comparison against the `Maximal` result comes from the returned values. +*) + +let droppedGroups = + inferred + |> List.filter (fun g -> + inferredMinimal + |> List.forall (fun m -> m.GroupOfProteinIDs <> g.GroupOfProteinIDs)) + |> List.map (fun g -> g.GroupOfProteinIDs) + +printfn "Maximal: %i groups, Minimal: %i groups" inferred.Length inferredMinimal.Length +printfn "dropped under Minimal: %s" (String.concat ", " droppedGroups) + +(** +```text +Maximal: 7 groups, Minimal: 6 groups +dropped under Minimal: Cre02.g095150.t1;Cre03.g144807.t1 +``` + +Under `Minimal` strictness the composite paralog group disappears, because +the two paralogs already explain the shared peptide on their own. The other +six groups come out identical under both settings. + +## Scoring the groups against reversed proteins + +A protein list needs the same error control as the peptide list before it, +again through a score compared against a decoy. The module +scores a protein group by summing the PSM scores of its peptides, and scores +its decoy counterpart from the peptides of the reversed protein sequences +that the search also matched. `PSMInput` is the row shape the module reads +from PSM result files: the peptide sequence and score of one match, plus +the database identifier of the peptide. Here the rows are built in memory, +one list per MS run, with deterministic scores on an Andromeda-like scale +where confident matches reach high double digits and random matches stay +low. +*) + +let psm id sequence score : ProteinInference.PSMInput = + { PepSequenceID = id; Seq = sequence; Score = score } + +let run1 = + [ + // Accepted target peptides. The one-hit wonder LISWYDNEWGYSNR + // scores weakly by design. + "VLNTWADIINR", 88.7 + "ETTENESANEGYR", 74.5 + "DTDILAAFR", 102.8 + "LTYYTPDYVVR", 68.9 + "SSYLDGSAPGDFGFDPLGLGK", 83.6 + "NLAGDVIGTR", 57.2 + "SYELPDGQVITIGNER", 79.4 + "VAPEEHPVLLTEAPLNPK", 88.1 + "AGFAGDDAPR", 61.7 + "LISWYDNEWGYSNR", 45.3 + // Decoy matches: some spectra matched peptides of the reversed + // protein sequences. Their scores sit in the random-match range. + "NIIDAWTNLVR", 18.3 + "VVYDPTYYTLR", 24.6 + "TGIVDGALNR", 21.4 + "ENGITIVQGDPLEYSR", 15.8 + "PNLPAETLLVPHEEPAVK", 27.5 + "NSYGWENDYWSILR", 26.7 + "GLTSAFVEK", 21.5 + ] + |> List.mapi (fun i (sequence, score) -> psm i sequence score) + +let run2 = + [ + // A second injection rescored two peptides. + "VLNTWADIINR", 96.2 + "DTDILAAFR", 95.1 + ] + |> List.mapi (fun i (sequence, score) -> psm (100 + i) sequence score) + +let peptideScores = ProteinInference.createPeptideScoreMap [run1; run2] + +printfn "best score for VLNTWADIINR: %.1f" peptideScores.["VLNTWADIINR"] + +(** +```text +best score for VLNTWADIINR: 96.2 +``` + +`createPeptideScoreMap` flattens the runs and keeps the best score seen for +each peptide sequence, so the rescored `VLNTWADIINR` carries 96.2 from the +second run while `DTDILAAFR` keeps its 102.8 from the first. + +The decoy side starts from the reverse digest: for every accession, the +peptides that its reversed sequence produces. The peptides follow the +reverse-and-keep-cleavage-site convention of common search engines, and the +score map above already contains the low scores they earned. +`createReverseProteinScores` sums, per reversed protein, the scores of those +of its peptides that were actually observed. The reversed one-hit wonder +picked up two such matches, and their sum is about to matter. +*) + +let reversedProteins = + [| + "psbA", [|"NIIDAWTNLVR"|] + "rbcL", [|"VVYDPTYYTLR"|] + // Both splice variants produce the same reversed peptide. + "Cre01.g001000.t1", [|"TGIVDGALNR"|] + "Cre01.g001000.t2", [|"TGIVDGALNR"|] + "Cre02.g095150.t1", [|"ENGITIVQGDPLEYSR"|] + "Cre03.g144807.t1", [|"PNLPAETLLVPHEEPAVK"|] + "Cre06.g250100.t1", [|"NSYGWENDYWSILR"; "GLTSAFVEK"|] + |] + +let decoyScores = ProteinInference.createReverseProteinScores reversedProteins peptideScores + +(** +With both maps in place, each inferred group receives its two scores. +`assignPeptideScores` sums the scores of the peptides the group kept for +quantification, and `assignDecoyScoreToTargetScore` takes the best decoy +score among the group's accessions. `createInferredProteinClassItemScored` +assembles the record the downstream estimators work with: these entries +describe target proteins found in the search database, and the +`DecoyHasBetterScore` flag records the picked target-decoy comparison of +the two scores. +*) + +let scored = + inferred + |> List.map (fun g -> + let targetScore = ProteinInference.assignPeptideScores g.PeptideSequence peptideScores + let decoyScore = ProteinInference.assignDecoyScoreToTargetScore g.GroupOfProteinIDs decoyScores + ProteinInference.createInferredProteinClassItemScored + g.GroupOfProteinIDs + g.Class + g.PeptideSequence + targetScore + decoyScore + false // these are target-side entries + (decoyScore > targetScore) // the picked target-decoy comparison + true) // every accession is in the search DB + |> Array.ofList + +for s in scored do + printfn "%-33s target %5.1f decoy %4.1f decoy wins %b" + s.GroupOfProteinIDs s.TargetScore s.DecoyScore s.DecoyHasBetterScore + +(** +```text +psbA target 170.7 decoy 18.3 decoy wins false +rbcL target 171.7 decoy 24.6 decoy wins false +Cre02.g095150.t1 target 79.4 decoy 15.8 decoy wins false +Cre03.g144807.t1 target 88.1 decoy 27.5 decoy wins false +Cre06.g250100.t1 target 45.3 decoy 48.2 decoy wins true +Cre01.g001000.t1;Cre01.g001000.t2 target 140.8 decoy 21.4 decoy wins false +Cre02.g095150.t1;Cre03.g144807.t1 target 61.7 decoy 27.5 decoy wins false +``` + +Six groups beat their decoy counterpart clearly. The one-hit wonder loses: +its single peptide scored 45.3 while the two random matches on its reversed +sequence sum to 48.2, so the group is flagged as a decoy win. This is the +protein-level analogue of a wrong PSM, and having such a case in the set is +what gives the error estimators of the next section something to measure. + +## Estimating the protein-level FDR + +`calculateFDRwithDecoyTargetRatio` is the direct estimate: the number of +decoy wins divided by the number of target wins, the protein-level version +of the decoy counting from the [FDR control]({{root}}04_01_fdr_control.html) +page. +*) + +let ratioFDR = ProteinInference.calculateFDRwithDecoyTargetRatio scored + +printfn "decoy/target ratio FDR: %.4f" ratioFDR + +(** +```text +decoy/target ratio FDR: 0.1667 +``` + +One decoy win against six target wins gives an estimated FDR of about 17 +percent. The number is high because the set is tiny, and it says that +roughly one of the six reported groups could be wrong at this acceptance. + +`calculateFDRwithMAYU` refines the estimate with the MAYU model introduced +in the MAYU section of the [FDR control]({{root}}04_01_fdr_control.html) +page. It needs the search database as accession and sequence pairs, because +the model stratifies proteins by sequence length before applying the +hypergeometric expectation per length bin, and unidentified database +proteins enter those bins too. Only the length of each sequence is used, so +placeholder residue strings of realistic lengths stand in for the real +sequences here. +*) + +let proteinsFromDB = + [| + // The seven accessions of the example, with realistic lengths. + "psbA", String.replicate 353 "A" + "rbcL", String.replicate 475 "A" + "Cre01.g001000.t1", String.replicate 254 "A" + "Cre01.g001000.t2", String.replicate 231 "A" + "Cre02.g095150.t1", String.replicate 380 "A" + "Cre03.g144807.t1", String.replicate 377 "A" + "Cre06.g250100.t1", String.replicate 335 "A" + // Database proteins that were never identified in this run. + "Cre09.g396920.t1", String.replicate 497 "A" + "Cre12.g554400.t1", String.replicate 155 "A" + "Cre16.g678000.t1", String.replicate 612 "A" + |] + +let mayuFDR = ProteinInference.calculateFDRwithMAYU scored proteinsFromDB + +printfn "MAYU FDR: %.4f" mayuFDR + +(** +```text +MAYU FDR: 0.1111 +``` + +MAYU lands somewhat below the raw ratio. It converts the decoy observation +in its length bin into an expected number of false positives among the +targets, and that expectation stays below one here, so the decoy win enters +the estimate with less than the full weight the ratio gave it. + +## Attaching protein q-values + +For the final list every group receives a q-value, computed from the same +score axis the estimators used. `FDRControl.calculateQValueStorey` from the +[FDR control]({{root}}04_01_fdr_control.html) page works on any record type +once it is told how to read the data, so it applies to the scored groups +directly. The decoy flag is `DecoyHasBetterScore`, and with the two score +selectors a decoy win enters the estimate at its decoy score while a +target win enters at its target score. The returned function +maps a score to its q-value, and `assignQValueToIPCIS` applies it to each +record, wrapping the result in an `InferredProteinClassItemQValue`. +*) + +let proteinQValue = + FDRControl.calculateQValueStorey + scored + (fun s -> s.DecoyHasBetterScore) + (fun s -> s.DecoyScore) + (fun s -> s.TargetScore) + +let qValued = + scored + |> Array.map (ProteinInference.assignQValueToIPCIS proteinQValue) + +for q in qValued do + printfn "%-33s score %5.1f q %6.4f" + q.InfProtClassItem.GroupOfProteinIDs + q.InfProtClassItem.TargetScore + q.QValue + +(** +```text +psbA score 170.7 q 0.0000 +rbcL score 171.7 q 0.0000 +Cre02.g095150.t1 score 79.4 q 0.0000 +Cre03.g144807.t1 score 88.1 q 0.0000 +Cre06.g250100.t1 score 45.3 q 0.2025 +Cre01.g001000.t1;Cre01.g001000.t2 score 140.8 q 0.0000 +Cre02.g095150.t1;Cre03.g144807.t1 score 61.7 q 0.0000 +``` + +Every group that beat its decoy counterpart sits above the score of the +single decoy win, so accepting any of them accumulates no decoy evidence +and their q-values are zero. The one-hit wonder sits at the bottom of the +score axis and collects the worst q-value of the run. Filtering this list +at the usual one percent threshold keeps the six confident groups and +removes the one-hit wonder. + +## Rendering the inference report with BioFSharp.Mz.Vis + +`BioFSharp.Mz.Vis` is the second library of this repository. It holds the +visual reporting that pipeline runs produce alongside their result tables, +built on Plotly.NET, and it ships as its own assembly so that the core +library carries no charting dependency. + +Its `ProteinInference.qValueHitsVisualization` renders the q-value results +of this page into an HTML report. The arguments are the histogram bandwidth +on the score axis, the array of q-value records, the output path stem and a +flag for grouped output. With `groupFiles = false` the function appends +`_QValueGraph` to the path stem and Plotly adds the `.html` extension. A +bandwidth of 10 suits the score axis of this example, which spans from the +forties up to about 170. +*) + +let reportStem = + System.IO.Path.Combine(System.IO.Path.GetTempPath(), "chlamy_protein_inference") + +BioFSharp.Mz.Vis.ProteinInference.qValueHitsVisualization 10.0 qValued reportStem false + +let reportPath = reportStem + "_QValueGraph.html" + +printfn "report exists: %b" (System.IO.File.Exists reportPath) +printfn "report size: %i KB" (System.IO.FileInfo(reportPath).Length / 1024L) + +(** +```text +report exists: true +report size: 11 KB +``` + +The report is a single Plotly figure. Column traces show the score +histograms of the target groups and of the decoy wins, drawn against a +relative frequency axis on the left and accompanied by an absolute count +axis on the right. A scatter trace on the same score axis draws each +group's q-value over its score, which on real data traces the rising error +curve toward low scores. In this small run the separation is easy to read: +the decoy-win bar sits at the low end of the score axis below every +confident target bar, and the q-value points are flat at zero everywhere +above it. The HTML loads plotly.js from a CDN, so viewing the report needs +network access. + +## The end of the identification pipeline + +The q-valued protein list is the final product of the identification side of the +pipeline. What remains is connecting the identities to amounts: the ion areas +determined by +[quantification]({{root}}03_01_quantification.html) are aggregated over exactly +these protein groups, using the peptides each group kept for quantification. +*)