diff --git a/README.md b/README.md
index 5dd7a5a..507e256 100644
--- a/README.md
+++ b/README.md
@@ -1,4 +1,4 @@
-## Getting Started
+## Getting Started
```sh
# Install hifiasm (requiring g++ and zlib)
@@ -19,23 +19,40 @@ hifiasm -o HG002.asm -t32 HG002-file1.fq.gz HG002-file2.fq.gz
yak count -b37 -t16 -o pat.yak <(cat pat_1.fq.gz pat_2.fq.gz) <(cat pat_1.fq.gz pat_2.fq.gz)
yak count -b37 -t16 -o mat.yak <(cat mat_1.fq.gz mat_2.fq.gz) <(cat mat_1.fq.gz mat_2.fq.gz)
hifiasm -o HG002.asm -t32 -1 pat.yak -2 mat.yak HG002-HiFi.fa.gz
+
+# Hi-C phasing with paired-end short reads in two FASTQ files
+hifiasm -o HG002.asm --h1 read1.fq.gz --h2 read2.fq.gz HG002-HiFi.fq.gz
```
-## Introduction
+## Table of Contents
-Hifiasm is a fast haplotype-resolved de novo assembler for PacBio Hifi reads.
-It can assemble a human genome in several hours and works with the California
-redwood genome, one of the most complex genomes sequenced so far. Hifiasm can
-produce primary/alternate assemblies of quality competitive with the best
-assemblers. It also introduces a new graph binning algorithm and achieves
-the best haplotype-resolved assembly given trio data.
+- [Getting Started](#started)
+- [Introduction](#intro)
+- [Why Hifiasm?](#why)
+- [Usage](#use)
+ - [Assembling HiFi reads without additional data types](#hifionly)
+ - [Hi-C integration](#hic)
+ - [Trio binning](#trio)
+ - [Output files](#output)
+- [Results](#results)
+- [Getting Help](#help)
+- [Limitations](#limit)
+- [Citing Hifiasm](#cite)
-## Why Hifiasm?
+## Introduction
+
+Hifiasm is a fast haplotype-resolved de novo assembler for PacBio HiFi reads.
+It can assemble a human genome in several hours and assemble a ~30Gb California
+redwood genome in a few days. Hifiasm emits partially phased assemblies of
+quality competitive with the best assemblers. Given parental short reads or
+Hi-C data, it produces arguably the best haplotype-resolved assemblies so far.
+
+## Why Hifiasm?
* Hifiasm delivers high-quality assemblies. It tends to generate longer contigs
and resolve more segmental duplications than other assemblers.
-* Given sequence reads from the parents, hifiasm can produce overall the best
+* Given Hi-C reads or short reads from the parents, hifiasm can produce overall the best
haplotype-resolved assembly so far. It is the assembler of choice by the
[Human Pangenome Project][hpp] for the first batch of samples.
@@ -47,13 +64,15 @@ the best haplotype-resolved assembly given trio data.
* Hifiasm is fast. It can assemble a human genome in half a day and assemble a
~30Gb redwood genome in three days. No genome is too large for hifiasm.
-* Hifiasm is trivial to install and easy to use. It does not required python,
- R or C++11 compilers and can be compiled into a single executable. The
+* Hifiasm is trivial to install and easy to use. It does not required Python,
+ R or C++11 compilers, and can be compiled into a single executable. The
default setting works well with a variety of genomes.
[hpp]: https://humanpangenome.org
-## Usage
+## Usage
+
+### Assembling HiFi reads without additional data types
A typical hifiasm command line looks like:
```sh
@@ -61,11 +80,21 @@ hifiasm -o NA12878.asm -t 32 NA12878.fq.gz
```
where `NA12878.fq.gz` provides the input reads, `-t` sets the number of CPUs in
use and `-o` specifies the prefix of output files. For this example, the
-primary contigs are written to `NA12878.asm.p_ctg.gfa` and alternate contigs to
-`NA12878.asm.a_ctg.gfa`. At the first run, hifiasm saves corrected reads and
+primary contigs are written to `NA12878.asm.bp.p_ctg.gfa` and alternate contigs to
+`NA12878.asm.bp.a_ctg.gfa`. Since v0.15, hifiasm also produces two sets of
+partially phased contigs at `NA12878.asm.bp.hap?.p_ctg.gfa`. This pair of files
+can be thought to represent the two haplotypes in a diploid genome, though with
+occasional switch errors. The frequency of switches is determined by the
+heterozygosity of the input sample.
+
+At the first run, hifiasm saves corrected reads and
overlaps to disk as `NA12878.asm.*.bin`. It reuses the saved results to avoid
the time-consuming all-vs-all overlap calculation next time. You may specify
`-i` to ignore precomputed overlaps and redo overlapping from raw reads.
+You can also dump error corrected in FASTA and/or overlaps in PAF with
+```sh
+hifiasm -o NA12878.asm -t 32 --write-paf --write-ec /dev/null
+```
Hifiasm purges haplotig duplications by default. For inbred or homozygous
genomes, you may disable purging with option `-l0`. Old HiFi reads may contain
@@ -75,7 +104,24 @@ bloom filter which takes 16GB memory at the beginning. For genomes much larger
than human, applying `-f38` or even `-f39` is preferred to save memory on k-mer
counting.
-When parental short reads are available, hifiasm can generate a pair of
+### Hi-C integration
+
+Hifiasm can generate a pair of haplotype-resolved assemblies with paired-end
+Hi-C reads:
+```sh
+hifiasm -o NA12878.asm -t32 --h1 read1.fq.gz --h2 read2.fq.gz HiFi-reads.fq.gz
+```
+In this mode, each contig is supposed to be a haplotig, which by definition
+comes from one parental haplotype only. Hifiasm often puts all contigs from the
+same parental chromosome in one assembly. It has cleanly separated chrX and
+chrY for a human male dataset. Nonetheless, phasing across centromeres is
+challenging. Users should not expect hifiasm to phase entire chromosomes at the
+moment. Also, contigs from different parental chromosomes are randomly mixed as
+it is just not possible to phase across chromosomes with Hi-C.
+
+### Trio binning
+
+When parental short reads are available, hifiasm can also generate a pair of
haplotype-resolved assemblies with trio binning. To perform such assembly, you
need to count k-mers first with [yak][yak] first and then do assembly:
```sh
@@ -85,19 +131,15 @@ hifiasm -o NA12878.asm -t 32 -1 pat.yak -2 mat.yak NA12878.fq.gz
```
Here `NA12878.asm.hap1.p_ctg.gfa` and `NA12878.asm.hap2.p_ctg.gfa` give the two
haplotype assemblies. In the binning mode, hifiasm does not purge haplotig
-duplications by default. Because hifiasm reuses saved overlaps, you can
+duplicates by default. Because hifiasm reuses saved overlaps, you can
generate both primary/alternate assemblies and trio binning assemblies with
```sh
hifiasm -o NA12878.asm -t 32 NA12878.fq.gz 2> NA12878.asm.pri.log
hifiasm -o NA12878.asm -t 32 -1 pat.yak -2 mat.yak /dev/null 2> NA12878.asm.trio.log
```
-The second command line will run much faster than the first. You can also dump
-error corrected in FASTA and/or overlaps in PAF with
-```sh
-hifiasm -o NA12878.asm -t 32 --write-paf --write-ec /dev/null
-```
+The second command line will run much faster than the first.
-## Output files
+### Output files
For non-trio assembly, hifiasm generates the following files:
@@ -126,9 +168,9 @@ For trio assembly, hifiasm generates the following files:
Hifiasm writes error corrected reads to the *prefix*.ec.bin binary file and
writes overlaps to *prefix*.ovlp.source.bin and *prefix*.ovlp.reverse.bin.
-## Results
+## Results
-The following table shows the statistics of several hifiasm primary assemblies:
+The following table shows the statistics of several hifiasm primary assemblies assembled with v0.12:
|Dataset|Size|Cov.|Asm options|CPU time|Wall time|RAM| N50|
|:---------------|-----:|-----:|:---------------------|-------:|--------:|----:|----------------:|
@@ -155,7 +197,10 @@ redwood genome in a few days on a single machine. For trio binning assembly:
|:---------------|-----:|-------:|--------:|----:|----------------:|
|[HG00733][HG00733-data], [\[father\]][HG00731-data], [\[mother\]][HG00732-data]|×33|269.1h|6.9h|135G|35.1Mb (paternal), 34.9Mb (maternal)|
|[HG002][NA24385-data], [\[father\]][NA24149-data], [\[mother\]][NA24143-data]|×36|305.4h|7.7h|137G|41.0Mb (paternal), 40.8Mb (maternal)|
+
+
[HG00733-data]: https://www.ebi.ac.uk/ena/data/view/ERX3831682
[HG00731-data]: https://www.ebi.ac.uk/ena/data/view/ERR3241754
@@ -167,33 +212,32 @@ redwood genome in a few days on a single machine. For trio binning assembly:
[NA12891-data]: https://www.ebi.ac.uk/ena/data/view/ERR194160
[NA12892-data]: https://www.ebi.ac.uk/ena/data/view/ERR194161
-Except NA12878, the assemblies above were produced by hifiasm v0.12 and can be
-downloaded at
-```txt
-ftp://ftp.dfci.harvard.edu/pub/hli/hifiasm/submission/hifiasm-0.12/
-```
-NA12878 was assembled with an older version of hifiasm and is available at
-```txt
-ftp://ftp.dfci.harvard.edu/pub/hli/hifiasm/NA12878-r253/
-```
-
+Human assemblies above can be acquired [from Zenodo][zenodo-human] and
+non-human ones are available [here][zenodo-nonh].
+[zenodo-human]: https://zenodo.org/record/4393631
+[zenodo-nonh]: https://zenodo.org/record/4393750
[unitig]: http://wgs-assembler.sourceforge.net/wiki/index.php/Celera_Assembler_Terminology
[gfa]: https://github.com/pmelsted/GFA-spec/blob/master/GFA-spec.md
[paf]: https://github.com/lh3/miniasm/blob/master/PAF.md
[yak]: https://github.com/lh3/yak
-## Getting Help
+## Getting Help
For detailed description of options, please see `man ./hifiasm.1`. The `-h`
option of hifiasm also provides brief description of options. If you have
further questions, please raise an issue at the [issue
page](https://github.com/chhylp123/hifiasm/issues).
-## Limitations
+## Limitations
1. Purging haplotig duplications may introduce misassemblies.
-## Citation
+## Citating Hifiasm
-Cheng, H., Concepcion, G.T., Feng, X., Zhang, H., Li H. Haplotype-resolved de novo assembly using phased assembly graphs with hifiasm. Nat Methods 18, 170–175 (2021). https://doi.org/10.1038/s41592-020-01056-5
+If you use hifiasm in your work, please cite:
+
+> Cheng, H., Concepcion, G.T., Feng, X., Zhang, H., Li H. (2021)
+> Haplotype-resolved de novo assembly using phased assembly graphs with
+> hifiasm. *Nat Methods*, **18**:170-175.
+> https://doi.org/10.1038/s41592-020-01056-5