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Alignment-free pangenome browser and analysis toolkit

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# Panagram: Interactive, alignment-free pan-genome browser

[![Release](https://img.shields.io/github/v/release/kjenike/panagram?label=Release)](https://github.com/kjenike/panagram/releases)
[![Integration](https://github.com/kjenike/panagram/actions/workflows/integration.yaml/badge.svg)](https://github.com/kjenike/panagram/actions/workflows/integration.yaml)

#### Katie Jenike, Nicole Brown, Sam Kovaka, Robin Burns, Shujun Ou, Stephen Hwang, Srividya Ramakrishnan, Ben Langmead, Elinor Karlsson, Zach Lippman, Ian R Henderson, Michael C Schatz

Welcome to Panagram! Panagram is
[an alignment-free pan-genome viewer](https://www.dropbox.com/s/g7snjgr8bs6c2uj/2023.01.17.Panagram.pdf).

# Installation

Requires python version >=3.11, pip, samtools, tabix, and bgzip from HTSlib. All other dependencies
should be automatically installed via pip.

Installation may fail if pip is not up-to-date or if setuptools is not up-to-date. In order to
update pip and setuptools run:

```bash
pip install --upgrade pip
pip install --upgrade setuptools
```

Once dependencies are installed, run:

```bash
git clone --recursive https://github.com/kjenike/panagram.git
cd panagram
pip install .
```

The `--recursive` option is required to install the KMC dependency. If you forget to include it, you
can update the repository with the command `git submodule update --init`.

## Notes

Panagram relies on [KMC](https://github.com/refresh-bio/KMC) to build its kmer index. This should be
installed automatically, however it is possible that the KMC installation will fail but panagram
will successfully install. In this case `panagram view` can be run, but `panagram index` will return
an error. You may be able to debug the KMC installation by running `make -C KMC py_kmc_api` and
attempting to fix any errors, then re-run `pip install -v .` after the errors are fixed. An
alternative fix is to manually update pybind11 in the KMC directory. You will need to replace the
"panagram/KMC/py_kmc_api/libs/pybind11/include" directory with the latest version at
<https://github.com/pybind/pybind11/tree/master/include/pybind11>. We are actively working to fix
this for a smoother installation.

# Running

Panagram runs in two steps, the anchoring step (index command) and viewing (view command).

## Anchoring

To prepare Panagram for anchoring, run:

```bash
panagram index <samples.tsv> -k <k> --prepare
```

To choose which genomes will act as anchors, use
`--anchor_genomes <one or more space-separated names>`. A common choice is to use one or more
reference genomes as anchors.

To run the indexing step, start by preparing the panagram index. It is best to create an empty
folder that will act as Panagram's index folder. Within this folder, create a subfolder called
FASTAS; this is where you can place any FASTAS that you want to include in your pan-genome. You can
also create a folder called GFFs; if you have any annotation files in GFF3 format, you can place
them in here. Next, you will need to tell Panagram where your FASTAS and GFFs are. For this, you
will need a tsv file with a list of the samples.

The samples.tsv file should contain one sample per line. You need the name, fasta, and gff columns.
On each line include the sample name and minimally the FASTA file location. Currently genome names
should only contain alphanumeric characters and underscores due to KMC requirements. If you have no
annotations for a file, leave the gff column blank. If you have multiple annotation files per
sample, you can concatenate them into one gff file. The id and anchor columns will be created by
Panagram. See below for an example.

```tsv
name	fasta	gff	id	anchor
sample1	FASTAS/sample1.fasta	GFFS/sample1.gff	0	True
sample2	FASTAS/sample2.fasta	GFFS/sample2.gff	1	True
sample3	FASTAS/sample3.fasta	GFFS/sample3.gff	2	True
sample4	FASTAS/sample4.fasta	GFFS/sample4.gff	3	True
sample5	FASTAS/sample5.fasta	GFFS/sample5.gff	4	True
sample6	FASTAS/sample6.fasta	GFFS/sample6.gff	5	True
```

As another example, this samples.tsv would be a comparison with just two genomes.

```tsv
name	fasta	gff	id	anchor
Col_0	FASTAS/wlod_Col-0.ragtag_scaffolds.fa	GFFS/wlod_Col-0.ragtag_scaffolds.gff	0	True
Tanz_1	FASTAS/wlod_Tanz-1.patch.scaffold.Chr.fa	GFFS/wlod_Tanz-1.patch.scaffold.Chr.gff	1	True
```

It is super important that any gff files are in the correct format. We strongly suggest that if you
run into any problems, you first check the format annotation format. This can be done with command
line tools like gff3validator or online at
[GenomeTools](https://genometools.org/cgi-bin/gff3validator.cgi).

Picking an acceptable kmer length for the data set can be tricky. For samples that are very
similar, a larger k may be more appropriate. While samples that are more diverged may benefit from
a smaller kmer length. The papers by
[Bonnie et al. (2024)](<https://www.cell.com/iscience/fulltext/S2589-0042(24)00275-X?uuid=uuid%3A8d061319-27f8-49ca-b7ee-0d33ec846225>)
and [Jenike et al. (2025)](https://pubmed.ncbi.nlm.nih.gov/39890468/) give some detail on picking
"good" kmer length, but if in doubt, k=21 usually works fine.

Once the preparation step is run, you can run Panagram's anchoring via snakemake and specify the
number of threads you want to use with this command:

```bash
snakemake --cores <num. threads> all
```

This step anchors KMC bitvectors to FASTA files to create a pan-kmer bitmap.

## View

Once anchoring is complete, navigate to the index folder and view your pan-genome with
`panagram view .` This runs a local Dash server. The pan-genome browser can be viewed at
<http://127.0.0.1:8050/> by default.

Here is the full set of flags you can choose:

```text
usage: panagram view [-h] <index_dir/> [genome] [chrom] [start] [end]
  index_dir           Panagram index directory
  genome              Initial anchor genome (optional)
  chrom               Initial chromosome (optional)
  start               Initial start coordinate (optional)
  end                 Initial end coordinate (optional)
  --ndebug            Run server in production mode (important for a public-
                      facing server)
  --port str          Server port (default: 8050)
  --host str          Server address (default: 127.0.0.1)
  --url_base str      A local URL prefix to use app-wide (passed to
                      Dash.dash(url_base_pathname=...)) (default: /)
  --bookmarks str     Bed file with bookmarked regions (default: None)
```

# Introgression Calling

Panagram's bitmap also enables calling introgressions between members of your pan-genome. For all
information on the introgression calling module, see the
[introgressions README](./panagram/introgressions).

# Bitdump

If you want to see the bitmap generated by Panagram, you can use the following:

```text
usage: panagram bitdump [-h] [-v bool] index_dir coords step
  Query pan-kmer bitmap generated by "panagram index"/

index_dir             Panagram index directory
  coords                Coordinates to query in chr:start-end format
  step                  Spacing between output kmers (optimized for multiples
                        of 100) (default: 1)
  -v bool, --verbose bool
                        Output the full bitmap (default: False)
```

# Example Run

First download the example_data.zip bacterial data from:
[http://data.schatz-lab.org/panagram/](http://data.schatz-lab.org/panagram/) or use this
[direct link](https://bx.bio.jhu.edu/data/panagram/example_data.zip).

Unzip the archive and you will find 5 bacterial genomes plus their annotations:

```bash
unzip example_data.zip
```

To run, first index the genomes:

```bash
cd example_data
panagram index samples.tsv -k 21 --prepare
snakemake --verbose --cores 30 all
```

Then you can panagram to visualize (from the example_data directory):

```bash
panagram view .
```

From there, you can view the results in your browser at
[http://127.0.0.1:8050/](http://127.0.0.1:8050).

# Hosting and Proxies

Panagram uses [Dash](https://dash.plotly.com/introduction) to serve the plotly visualizations. By
default the dedicated webserver runs on localhost (127.0.0.1) on port 8050, but you can reverse
proxy to a different port and path using a web engine such as [nginx](https://www.nginx.com/).

For nginx, first reconfigure your nginx configuration file to add (note to be very careful with the
use of the slash ('/') character):

```text
    location /panagram {
      proxy_pass http://127.0.0.1:8050;
    }
```

The restart nginx with:

```bash
systemctl stop nginx
systemctl start nginx
```

For a secure public-facing server, be sure to run with the option `panagram view --ndebug` to
disable debug mode. You may also wish to change the base URL path with the `--url_base` option, for
example to something like `--url_base /panagram/`. The port and host name can be specified by the
`--port` and `--host` options.

Finally you will need to run panagram using `panagram view <dir>`. You will probably want to run
this in a loop in case it needs to be restarted, such as:

```bash
until panagram view --ndebug .; do echo "restarting"; sleep 1; done
```

# Contributing and Support

Have a question or found a bug? [Open an issue](https://github.com/kjenike/panagram/issues)!

Like to contribute? Send us a pull request! We use the following tools for code quality and formatting:

- **ruff** for Python linting
- **mypy** for type checking
- **markdownlint-cli2** for Markdown linting
- **prettier** for Markdown formatting

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