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Table 1 Summary of algorithms and features of the examined read alignment methods. We surveyed 107 alignment tools published from 1988 to 2020 (indicated in column “Year of publication”). The table is sorted by year of publication, and then grouped according to the area(s) of application (indicated in column “Application”) within each year. In column “Indexing,” we document the algorithms used to index the genome (the first step in read alignment). In column “Global Positioning,” we document the algorithms used to determine a global position of the read in the reference genome (the second step). In column “Pairwise alignment,” we document the algorithm used to determine the similarity between the read and the corresponding region of the reference genome (the last step). SW, NW, HD, and DP stand for Smith-Waterman algorithm, Needleman-Wunsch algorithm, Hamming distance, and dynamic programming, respectively. In column “Wrapper,” we document the read alignment algorithms that are built on top of other read alignment tools. Finally, we report the maximum read length tested in the corresponding paper in column “Max. Read Length Tested in the Paper (bp).” The tested read length in each paper is not necessarily the maximum read length that each tool can handle

From: Technology dictates algorithms: recent developments in read alignment

Aligner

URL

Year of publication

Application

Indexing

Global Positioning

Pairwise alignment

Wrapper

Max. read length tested in the paper (bp)

Fix length seed

Spaced seed

Seed chaining

FASTA [30]

https://fasta.bioch.virginia.edu/fasta_www2/fasta_list2.shtml

1988

DNA

Hashing

Y

N

Y

SW and NW

N

1500

BLAST [31]

https://blast.ncbi.nlm.nih.gov/Blast.cgi

1990

DNA

Hashing

Y

N

Y

Non-DP Heuristic

N

73360

Gapped BLAST [32]

https://blast.ncbi.nlm.nih.gov/Blast.cgi

1997

DNA

Hashing

Y

N

Y

SW

N

246

SSAHA [33]

https://www.sanger.ac.uk/science/tools/ssaha

2001

DNA

Hashing

Y

N

N

NW

N

500

PatternHunter [34,35,36,37]

https://www.bioinfor.com/

2002

DNA

Hashing

Y

Y

Y

Non-DP heuristic

N

500

BLAT [38]

https://genome.ucsc.edu/cgi-bin/hgBlat

2002

DNA

Hashing

Y

N

Y

Non-DP heuristic

N

500

BLASTZ [39]

https://www.bx.psu.edu/miller_lab/

2003

DNA

Hashing

Y

N

N

SW

Y

3000

C4 [40]

https://github.com/nathanweeks/exonerate

2005

DNA

Hashing

Y

N

Y

Sparse DP

N

N/A

GMAP [41]

https://github.com/juliangehring/GMAP-GSNAP

2005

DNA

Hashing

N

N

Y

NW

N

N/A

BWT-SW [42]

https://github.com/mruffalo/bwt-sw

2008

DNA

BWT

Y

N

N

SW

N

2000

MAQ [43]

http://maq.sourceforge.net/maq-man.shtml

2008

DNA

Hashing

Y

Y

N

SW

N

63

RMAP [44]

https://github.com/smithlabcode/rmap

2008

DNA

Hashing

Y

N

N

HD

N

36

SOAP [45]

https://github.com/ShujiaHuang/SOAPaligner

2008

DNA

Hashing

Y

N

N

Non-DP heuristic

N

50

SOCS [46]

http://socs.biology.gatech.edu/

2008

DNA

Hashing

Y

N

N

Rabin-Karp Algorithm

N

35

SeqMap [47]

http://www-personal.umich.edu/~jianghui/seqmap/

2008

DNA

Hashing

Y

N

N

Non-DP Heuristic

N

30

ZOOM [48]

http://www.bioinfor.com/zoom-1-3-gui-release-next-gen-seq/

2008

DNA

Hashing

Y

Y

N

SW

N

36

QPALMA [49, 50]

http://www.raetschlab.org/suppl/qpalma

2008

RNA-Seq

Suffix array

Y

N

Y

SW

Y

36

BRAT [51]

http://compbio.cs.ucr.edu/brat/

2009

BS-Seq

Hashing

Y

N

N

HD

N

26

BSMAP [52]

https://github.com/genome-vendor/bsmap

2009

BS-Seq

Hashing

Y

N

N

HD

N

32

BFAST [53]

https://github.com/nh13/BFAST/

2009

DNA

Hashing

N

Y

N

SW

N

55

BWA [54]

https://github.com/lh3/bwa

2009

DNA

BWT-FM

N

N

N

Semi-Global

N

125

Bowtie [55]

http://bowtie-bio.sourceforge.net/manual.shtml

2009

DNA

BWT-FM

Y

N

N

HD

N

76

CloudBurst [56]

https://sourceforge.net/projects/cloudburst-bio/

2009

DNA

Hashing

Y

N

N

Landau-Vishkin

N

36

GNUMAP [57]

https://github.com/byucsl/gnumap

2009

DNA

Hashing

Y

N

Y

NW

N

36

GenomeMapper [58]

http://1001genomes.org/software/genomemapper_singleref.html

2009

DNA

Hashing

Y

N

Y

NW

N

200

MOM [59]

https://github.com/hugheaves/MOM

2009

DNA

Hashing

Y

N

N

HD

N

40

PASS [60]

http://pass.cribi.unipd.it/cgi-bin/pass.pl

2009

DNA

Hashing

Y

N

Y

NW

N

32

PerM [61]

https://code.google.com/archive/p/perm/downloads

2009

DNA

Hashing

Y

Y

N

HD

N

47

RazerS [62]

https://github.com/seqan/seqan/tree/master/apps/razers

2009

DNA

Hashing

Y

Y

Y

Myers Bit Vector

N

76

SHRiMP [63]

http://compbio.cs.toronto.edu/shrimp/

2009

DNA

Hashing

N

N

N

SW

N

35

SOAP2 [64]

https://github.com/ShujiaHuang/SOAPaligner

2009

DNA

BWT-FM

Y

N

N

SW

N

44

Slider [65]

http://www.bcgsc.ca/platform/bioinfo/software/slider

2009

DNA

Hashing

Y

N

N

HD

N

36

segemehl [66]

https://www.bioinf.uni-leipzig.de/Software/segemehl/

2009

DNA

Suffix array

N

N

Y

SW

N

35

TopHat [67]

https://ccb.jhu.edu/software/tophat/index.shtml

2009

RNA-Seq

BWT-FM

Y

N

N

HD

Y

42

BS-Seeker [68]

http://pellegrini-legacy.mcdb.ucla.edu/bs_seeker/BS_Seeker.html

2010

BS-Seq

BWT-FM

Y

N

N

HD

Y

36

BWA-SW [54]

https://github.com/lh3/bwa

2010

DNA

BWT-FM

N

N

N

SW

N

10000

GASSST [35]

http://www.irisa.fr/symbiose/projects/gassst/

2010

DNA

Hashing

Y

Y

Y

Semi-Global

N

500

GSNAP [37]

https://github.com/juliangehring/GMAP-GSNAP

2010

DNA

Hashing

Y

N

Y

Non-DP Heuristic

N

100

SMALT [69]

https://github.com/rcallahan/smalt

2010

DNA

Hashing

Y

N

Y

SW

N

150

Slider II [70]

http://www.bcgsc.ca/platform/bioinfo/software/SliderII

2010

DNA

Hashing

Y

N

N

HD

Y

42

VMATCH [71]

http://www.vmatch.de/

2010

DNA

Suffix array

Y

N

Y

SW

Y

N/A

mrsFAST [72]

https://github.com/sfu-compbio/mrsfast

2010

DNA

Hashing

Y

N

N

HD

N

100

MapSplice [73]

https://github.com/LiuBioinfo/MapSplice

2010

RNA-Seq

BWT-FM

Y

N

N

HD

Y

100

MicroRazerS [74]

https://github.com/seqan/seqan/tree/master/apps/micro_razers

2010

RNA-Seq

Hashing

Y

N

Y

HD

N

36

SpliceMap [75]

http://web.stanford.edu/group/wonglab/SpliceMap/

2010

RNA-Seq

Hashing

Y

N

N

HD

Y

50

Supersplat [76]

http://mocklerlab.org/tools/1/manual

2010

RNA-Seq

Hashing

N

N

N

NA

N

36

Bismark [77]

https://github.com/FelixKrueger/Bismark

2011

BS-Seq

BWT-FM

Y

N

Y

SW & NW

Y

50

LAST [78]

http://last.cbrc.jp/

2011

DNA/BS-Seq/RNA

Suffix array

N

Y

N

SW & NW

N

105

DynMap [79]

https://dl.acm.org/citation.cfm?id=2147845&dl=ACM&coll=DL

2011

DNA

Hashing

Y

N

N

NW

N

52

SHRiMP2 [80]

http://compbio.cs.toronto.edu/shrimp/

2011

DNA

Hashing

Y

Y

Y

SW

N

75

SNAP [81]

http://snap.cs.berkeley.edu/

2011

DNA

Hashing

Y

N

N

NW

N

10000

Stampy [82]

https://www.well.ox.ac.uk/project-stampy

2011

DNA

Hashing

Y

N

N

NW

N

4500


TMAP

https://github.com/iontorrent/TS/tree/master/Analysis/TMAP

2011

DNA

BWT-FM

N

N

Y

SW

N

N/A

X-Mate [83]

http://grimmond.imb.uq.edu.au/X-MATE/

2011

DNA

Hashing

N

N

N

Non-DP Heuristic

N

50

SOAPSplice [84]

http://soap.genomics.org.cn/soapsplice.html

2011

RNA-Seq

BWT-FM

Y

N

N

Non-DP Heuristic

N

150

BRAT-BW [51]

http://compbio.cs.ucr.edu/brat/

2012

BS-Seq

BWT-FM

N

N

N

HD

N

62

BLASR [85]

https://github.com/mchaisso/blasr/

2012

DNA

Suffix array

Y

N

Y

NW

N

8000

Batmis [86]

https://code.google.com/archive/p/batmis/

2012

DNA

BWT-ST

Y

N

N

HD

N

100

Bowtie2 [87]

http://bowtie-bio.sourceforge.net/bowtie2

2012

DNA

BWT-FM

Y

N

Y

SW & NW

N

400

GEM [88]

https://github.com/smarco/gem3-mapper

2012

DNA

BWT-FM

N

N

Y

SW & NW

N

150

RazerS3 [89]

https://github.com/seqan/seqan/tree/master/apps/razers3

2012

DNA

Hashing

Y

Y

Y

Banded Myers Bit Vector

N

800

SeqAlto [90]

https://web.stanford.edu/group/wonglab/seqalto/

2012

DNA

Hashing

Y

N

N

NW

N

200

SplazerS [91]

https://github.com/seqan/seqan/blob/master/apps/splazers/README

2012

DNA

Hashing

Y

N

Y

Banded Myers Bit Vector

N

150

WHAM [92]

http://pages.cs.wisc.edu/~jignesh/wham/

2012

DNA

Hashing

Y

N

N

NW

N

74

YAHA [93]

https://github.com/GregoryFaust/yaha

2012

DNA

Hashing

Y

N

Y

SW

N

10000

OSA [94]

http://www.arrayserver.com/wiki/index.php?title=OSA

2012

RNA-Seq

Hashing

Y

N

N

NA

N

100

Passion [95]

https://trac.nbic.nl/passion/

2012

RNA-Seq

Hashing

Y

N

Y

SW

Y

75

BS-Seeker2 [96]

https://github.com/BSSeeker/BSseeker2

2013

BS-Seq

BWT-FM

Y

N

Y

SW & NW

Y

250

Subread [97]

http://subread.sourceforge.net/

2013

DNA/RNA-Seq

Hashing

Y

Y

Y

SW

N

202

BWA-MEM [98]

https://github.com/lh3/bwa

2013

DNA

BWT-FM

N

N

Y

SW & NW

N

650

Masai [99]

http://www.seqan.de/projects/masai

2013

DNA

Suffix tree

N

N

Y

Banded Myers Bit Vector

N

150

NextGenMap [100]

http://cibiv.github.io/NextGenMap/

2013

DNA

Hashing

Y

N

N

SW & NW

N

250

SRmapper [101]

http://www.umsl.edu/~wongch/software.html

2013

DNA

Hashing

Y

N

N

HD

N

100

mrFAST [102]

https://github.com/BilkentCompGen/mrfast

2013

DNA

Hashing

Y

N

N

Semi-Global

N

180

CRAC [103]

http://crac.gforge.inria.fr/

2013

RNA-Seq

BWT-FM

Y

N

N

Non-DP Heuristic

N

200

STAR [104]

https://github.com/alexdobin/STAR

2013

RNA-Seq

Suffix array

N

N

Y

SW

N

5000

TopHat2 [105]

https://ccb.jhu.edu/software/tophat/index.shtml

2013

RNA-Seq

BWT-FM

Y

N

Y

SW & NW

Y

101

Subjunc [106]

http://subread.sourceforge.net/

2013

RNA-seq

Hashing

Y

Y

Y

NW

N

202

BWA-PSSM [107]

http://bwa-pssm.binf.ku.dk/

2014

DNA

BWT-FM

Y

N

N

SW

Y

100

CUSHAW3 [108]

http://cushaw3.sourceforge.net/homepage.htm#latest

2014

DNA

BWT-FM

Y

N

Y

SW & Semi-Global

N

100

Hobbes2 [109]

https://hobbes.ics.uci.edu/download.shtml

2014

DNA

Hashing

Y

N

Y

Banded Myers Bit Vector

N

100

MOSAIK [110]

https://github.com/wanpinglee/MOSAIK

2014

DNA

Hashing

Y

N

N

SW

N

100

hpg-Aligner [111]

https://github.com/opencb/hpg-aligner

2014

DNA

Suffix array

N

N

Y

SW

N

5000

mrsFAST-Ultra [112]

https://github.com/sfu-compbio/mrsfast

2014

DNA

Hashing

Y

N

N

HD

N

100

JAGuaR [113]

http://www.bcgsc.ca/platform/bioinfo/software/jaguar

2014

RNA-Seq

BWT-FM

Y

N

N

SW

Y

100

ContextMap 2 [114]

http://www.bio.ifi.lmu.de/ContextMap

2015

RNA-Seq

BWT-FM

Y

N

Y

SW & NW

Y

76

HISAT [115]

http://www.ccb.jhu.edu/software/hisat/index.shtml

2015

RNA-Seq

BWT-FM

Y

N

N

Non-DP Heuristic

N

100

ERNE 2 [116]

http://erne.sourceforge.net/

2016

DNA/BS-Seq

BWT-FM + hashing

Y

N

N

HD

N

100

GraphMap [117]

https://github.com/isovic/graphmap

2016

DNA

Hashing

Y

Y

Y

Semi-global

N

9000

NanoBLASTer [118]

https://github.com/ruhulsbu/NanoBLASTer

2016

DNA

Hashing

Y

N

Y

NW

N

7040

minimap [119]

https://github.com/lh3/minimap

2016

DNA

Hashing

Y

N

N

N/A

N

13000

rHAT [120]

https://github.com/dfguan/rHAT

2016

DNA

Hashing

Y

N

Y

SW

N

8000

KART [121]

https://github.com/hsinnan75/KART

2017

DNA

BWT-FM

N

N

Y

NW

N

7118

LAMSA [122]

https://github.com/hitbc/LAMSA

2017

DNA

BWT-FM + hashing

Y

N

Y

Sparse DP

Y

100000

DART [123]

https://github.com/hsinnan75/DART

2017

RNA-Seq

BWT-FM

N

N

Y

NW

N

251

minimap2 [124]

https://github.com/lh3/minimap2

2018

DNA/RNA-Seq

Hashing

Y

N

Y

NW

N

11628

DREAM-Yara [125]

https://gitlab.com/pirovc/dream_yara/

2018

DNA

BWT-FM

Y

N

N

Banded Myers Bit Vector

Y

150

MUMmer4 [126]

https://github.com/mummer4/mummer

2018

DNA

Suffix array

Y

N

Y

SW

Y

7821

NGMLR [127]

https://github.com/philres/ngmlr

2018

DNA

Hashing

Y

N

Y

SW

N

50000

lordFAST [128]

https://github.com/vpc-ccg/lordfast

2018

DNA

BWT-FM + hashing

N

N

Y

SW & NW

N

35489

BatMeth2 [129]

https://github.com/GuoliangLi-HZAU/BatMeth2/

2019

BS-Seq

BWT-FM

Y

N

Y

SW & NW

N

125

GraphMap2 [130]

https://github.com/lbcb-sci/graphmap2

2019

DNA/RNA-Seq

Hashing

Y

Y

Y

Semi-global

N

9000

Magic-BLAST [131]

https://github.com/ncbi/magicblast

2019

DNA/RNA-Seq

Hashing

Y

N

N

Non-DP Heuristic

N

90000

BWA-MEM2 [132]

https://github.com/bwa-mem2/bwa-mem2

2019

DNA

BWT-FM

N

N

Y

SW

N

650

HISAT2 [133]

https://ccb.jhu.edu/software/hisat2/index.shtml

2019

DNA

BWT-FM

Y

N

N

Non-DP Heuristic

N

100

deSALT [134]

https://github.com/hitbc/deSALT

2019

RNA-seq

Hashing

Y

N

Y

SW

N

8000

conLSH [135]

https://www.dropbox.com/s/3jcu4i240kyu2tc/source%20code%20conLSH_bio.tar.gz?dl=0

2020

DNA

Hashing

Y

N

Y

Sparse DP

N

8000