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Genomic Perspectives on Evolution in Bracken Fern
Joshua DerDepartment of Biology
PhD Defense, 27 May 2010
1Friday, May 28, 2010
Collage by Kristal Watrous
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
2Friday, May 28, 2010
Collage by Kristal Watrous
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
They all have complete sequenced genomes!
2Friday, May 28, 2010
Fruit fly Mouse
Model organisms (usually with small genomes)
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
3Friday, May 28, 2010
Yeast Grapes
Economically important species
Wine
+ =
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
4Friday, May 28, 2010
Rice Humans
Economically important species
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
5Friday, May 28, 2010
Platypus Lycophyte
Important evolutionary lineages
Sea Squirt
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
6Friday, May 28, 2010
Genomics in model organisms
• Generally have small genomes
• Decipher gene and genome function
• Understand networks of molecular interactions (i.e. systems biology)
• Translate new discoveries to benefit people (e.g. crop improvement and human health)
7Friday, May 28, 2010
The age of genomics
0
375
750
1125
1500
1995 1997 1999 2001 2003 2005 2007 2009
Number of complete genome sequences in Genbank, Feb. 2010(Data from Genomes OnLine Database v.3.0: www.genomesonline.org)
8Friday, May 28, 2010
“Next-generation” sequencing revolution
• Massive throughput
• Avoid sequence bias and labor associated with molecular cloning
• Huge reduction in cost per base
• Target sequencing for information-rich templates (e.g. gene space and transcriptomes)
9Friday, May 28, 2010
Next-generation platforms
• Second-generation (in vitro clonal amplification)
• Illumina
• SOLiD
• Roche 454
• Third-generation (single molecule sequencing)
• Helicos
• Pacific Biosciences
• Oxford Nanopore
10Friday, May 28, 2010
Genome-scale analyses can be performed on ANY organism
What’s your favorite organism?
11Friday, May 28, 2010
Genome-scale analyses can be performed on ANY organism
Woolly Mammoth
• Genome sequenced 2008
• Roche 454
Credit: Mauricio Anton
12Friday, May 28, 2010
Genome-scale analyses can be performed on ANY organism
Giant Panda
• Genome sequenced 2010
• Illumina
Photo: Aaron Logan
13Friday, May 28, 2010
Genome-scale analyses can be performed on ANY organism
• 1000 Genomes (human)
• 1001 Genomes (Arabidopsis)
• 1000 Plant and Animal Reference Genomes Project
• Genome 10K (vertebrates)
• OneKP (plant transcriptomes)
Let’s just sequence it all!
Photo: Natalie Maynor
14Friday, May 28, 2010
Reality check!
• We have limited resources
• Current technology still has limits
• Instrument capacity
• Throughput
• Read lengths
15Friday, May 28, 2010
So, how do we choose?
• Plants, of course!
• Two papers give us some guidance
16Friday, May 28, 2010
So, how do we choose?
• Choice must be considered within a phylogenetic framework
• Once we choose:
Do we need the full genome (yet)?
17Friday, May 28, 2010
Among plants, what is the last major (i.e. large and diverse) lineage
unsampled in a genome-scale project?
Ferns!
Photo: Robbin Moran
18Friday, May 28, 2010
Fern evolution
• Sister to seed plants
• Ancient lineage (Devonian)
• ~9000 extant species
• Geographic and ecological diversity
• Evolved and maintain independent gametophyte and sporophyte generations
Ferns
Lycophytes
Bryophytes
Seed Plants
19Friday, May 28, 2010
Fern evolution
• Sister to seed plants
• Ancient lineage (Devonian)
• ~9000 extant species
• Geographic and ecological diversity
• Evolved and maintain independent gametophyte and sporophyte generations
haploid spores (n)
meiosis
sperm (n)
egg (n)
zygote (2n)
Fern life cycle
syngamysporophyte (2n)
gametophyte (n)
20Friday, May 28, 2010
Challenges in fern genomics
• Limited agronomic importance
• Large genome sizes (avg. 10 Gb)
• High chromosome numbers (avg. n = 57)
• Extensive history of hybridization and polyploidy
Photo: Mike Windham
21Friday, May 28, 2010
Bracken fern: Pteridium
• Worldwide distribution
• Toxic to livestock and weedy in pasture
• Highly adaptable and phenotypically plastic
• Established culture techniques
• Paleopolyploid with diploid gene expression
• Genome size: 1C = 9.8 GbLindman. 1917-1926. Bilder ur Nordens Flora-508
22Friday, May 28, 2010
Bracken fern: Pteridium
Model system in the study of:
• Fern lifecycle
• Apomixis, apogamy, apospory
• Gametophyte development
• Pheromonal sex determination
• Cyanogenesis/Carcinogenesis
• Invasion ecology
• Climate change
Photo: John Thomson
23Friday, May 28, 2010
Dissertation outline1. Global phylogeny and biogeography of bracken
• Establish an evolutionary framework
2. Characterization of the bracken gametophyte transcriptome
• Leverage 454 sequencing to sample the expressed component of the huge nuclear genome
3. Chloroplast genome and RNA editing in bracken
• Combine genomic and expressed 454 sequence data to study chloroplast RNA metabolism
24Friday, May 28, 2010
1. Global phylogeny and biogeography of bracken
Citation: Joshua P. Der, John A. Thomson, Jeran K. Stratford and Paul G. Wolf. 2009. Global chloroplast phylogeny and biogeography of bracken (Pteridium: Dennstaedtiaceae). American Journal of Botany 96 (5): 1441-1449.
25Friday, May 28, 2010
Taxonomic instability• High phenotypic plasticity with
environmental conditions
• Few diagnostic morphological characters recognized
• Intermediate forms between geographic morphotypes
• Hybrid tetraploid species
• Regionally biased studiesThomson JA, Mickel JT, Mehltreter K. Botanical Journal of the Linnean Society, 2008, 157, 1–17.
Over 135 named forms!
26Friday, May 28, 2010
Phylogenetic objectives:
1. Establish a phylogenetic framework for the genus worldwide
2. Examine patterns of global biogeography
27Friday, May 28, 2010
Phylogeny
• 77 specimens sampled globally
• trnS-rps4 spacer+gene &rpl16 intron
• 43 parsimony informative sites
• 3 indel haplotypes detected
28Friday, May 28, 2010
Phylogeny
• 77 specimens sampled globally
• trnS-rps4 spacer+gene &rpl16 intron
• 43 parsimony informative sites
• 3 indel haplotypes detected
AC
B
28Friday, May 28, 2010
Biogeography
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Biogeography
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Biogeography
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30Friday, May 28, 2010
Phylogenetic conclusions
1. First global phylogenetic analysis in bracken
2. Biogeographic patterns are consistent with paleoclimatic events
3. This work contributes to a revision of the genus
31Friday, May 28, 2010
2. Gametophyte transcriptome
Coauthors: Michael S. Barker, Norman J. Wickett, Claude W. dePamphilis, Paul G. Wolf
32Friday, May 28, 2010
Transcriptome objectives
1. Develop an extensive expressed sequence resource in ferns
2. Glimpse into the nuclear genome of bracken
3. Characterize the transcriptome & identify active genes in the gametophyte generation
33Friday, May 28, 2010
Roche 454 sequence reads
Normalized cDNA derived from mature gametophytes
Reads were quality and length filtered, adapter and polyA/T trimmed
• Cleaned reads:! 681,722
• Mean length:!! 372.60 bp
• Total bases:! ! 254 Mb
Histogram of cleaned reads
Cleaned read length, maximum = 624
Num
ber o
f seq
uenc
es
0 100 200 300 400 500 600
050
0010
000
1500
0
34Friday, May 28, 2010
Transcriptome assembly
Histogram of transcriptome unigenes (CAP3)
Unigene length, largest transcript = 4897 bp
Num
ber o
f seq
uenc
es
0 500 1000 1500 2000 25000
2000
4000
6000
Total unigenes = 38889
Mean length = 685.76 bp
Total bases = 26.67 Mp
Unigenes
Number 38,889
Mean length (bp) 685.76
Mode (bp) 476
Range in length (bp) 86 - 4,897
Total bases (Mbp) 26.67
35Friday, May 28, 2010
Transcriptome coverage
Wall et. al., 2009. BMC Genomics 10:347
ESTcalc (simulation based model)ESTcalc (simulation based model)
Percent of the unique bases in the transcriptome: 87%
Percent of the genes with at least one read: 100%
19
338
Ultra conserved orthologs
8
155
Shared single copy genes
95%
36Friday, May 28, 2010
Transcriptome coverage
95% of unigenes captured after 403K reads sampled
1,357 reads to capture the last 10 unigenes
37Friday, May 28, 2010
Functional annotation
Localization of genes is predominantly in the nucleus, mitochondria, and plastids
cellular_component Level 5
endoplasmic reticulum
(317)
nucleoplasm (376)
vacuole (274)
Golgi apparatus
(212)
microbody (119)
plastid (3,613)
cytoskeleton (238)
nucleus (1,325)
endosome (10)
nucleolus (197)
nuclear lumen (555)
cytosol (448)
mitochondrion (1,967)
GO category: Cellular Component
38Friday, May 28, 2010
Functional annotation
Two main biological processes involve metabolism and cellular machinery
GO category: Biological Processbiological_process Level 2
multicellular organismal
process (166)
localization (1,713)
multi-organism process (15)
growth (41)
establishment of localization
(1,713)
reproduction (73)
biological regulation (853)
developmental process (194)
reproductive process (29)
cellular process (7,432)
regulation of biological
process (716)
response to stimulus (908)
metabolic process (7,641)
39Friday, May 28, 2010
Functional annotation
Two main molecular functions are binding (DNA, RNA, and protein) and catalytic activity (hydrolase and transferase activity)
GO category: Molecular functionmolecular_function Level 2
enzyme regulator
activity (106)
binding (8,120)
transcription regulator
activity (409)
structural molecule
activity (542)
translation regulator activity (1)
transporter activity (908)
molecular transducer
activity (357)
catalytic activity (7,915)
40Friday, May 28, 2010
Comparative genomics
Of 38,889 unigenes, 24,897 had a positive blastx hit
• 23,152 in Arabidopsis
• 22,474 in Physcomitrella
• 16,352 in Selaginella
41Friday, May 28, 2010
Transcriptome conclusions
1. First comprehensive analysis of gene expression in a fern
2. Most extensive sequence resource available in ferns
42Friday, May 28, 2010
3. RNA editing in the chloroplast genome
Coauthors: Aaron M. Duffy, Matt Kusner, Chen Gu, Paul Overvoorde, Paul G. Wolf
43Friday, May 28, 2010
RNA editing in plants• Conversion of C to U or U
to C nucleotide in RNA molecules relative to the encoding DNA
• More abundant in mitochondria than chloroplasts or nuclear genomes
• Most abundant in seed-free vascular plants and hornworts
Ferns (300; ?)
Lycophytes (?; 1450)
Mosses (1; 11)
Seed Plants (30; 500)
Hornworts (900; 600)
Leafy liverworts (?; 600)Thalloid liverwort (none)
Lineage (cp; mt)
44Friday, May 28, 2010
Chloroplast genome objectives:
1. Reconstruct the complete chloroplast genome from total genomic Roche 454 data
2. Use expressed transcript data to identify RNA editing sites, de novo
45Friday, May 28, 2010
Chloroplast genomeGenomic 454 read lengths
Read length (maximum = 1363)
Num
ber o
f rea
ds
0 100 200 300 400 500 600 700
050
0015
000
2500
035
000
711,178 reads216.19 Mbp
152,362 bp34.5x coverage
46Friday, May 28, 2010
Genome annotation
Ribosomal RNAs 4Transfer RNAs 29Photosystem I 5Photosystem II 15Cytochrome 6ATP synthase 6Rubisco (rbcL) 1Chlorophyll biosynthesis 3NADH dehydrogenase 11Ribosomal proteins 22RNA polymerase 4Intron maturase (matK) 1Miscellaneous proteins 5Hypothetical proteins 5
117 total genes
47Friday, May 28, 2010
RNA editing
Percent chloroplast transcripts 2.8%Percent of chloroplast genome 80.67%Percent of chloroplast sites edited 0.73%Total RNA editing sites detected 851
C to U RNA editing 551U to C RNA editing 300RNA editing in the Inverted Repeat 233
RNA editing in protein coding sequence 660Modified start codons 26Premature stop codon repair 37Repair proper stop codon 6
RNA editing in tRNA genes 15RNA editing in rRNA genes 168RNA editing in introns 12
48Friday, May 28, 2010
RNA editing
49Friday, May 28, 2010
Chloroplast genome conclusions
1. Utilized a novel method to reconstruct the chloroplast genome
• Pteridium aquilinum has the chloroplast gene set and gene order of Adiantum capillis-veneris
2. First to use deep RNA sequencing to identify RNA editing in chloroplasts
• Identified a large number of RNA editing sites in the chloroplast genome
50Friday, May 28, 2010
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
51Friday, May 28, 2010
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
Conclusions1. Next generation sequencing has enabled
genome-scale studies in non-model organisms
2. Large data sets present new challenges for analysis
3. Incredible opportunities for novel studies and analyses in new organisms
Friday, May 28, 2010
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Future work
1. Extract mitochondrial genome sequences
2. Sequence the sporophyte transcriptome
3. Explore gene space and transposable elements in the nuclear genome
Collage by Kristal Watrous
52Friday, May 28, 2010
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
AcknowledgmentsPhD Committee: Coauthors: Claude dePamphilisPaul Wolf Mike Pfrender Jeran Stratford Aaron DuffyCarol VonDohlen Karen Mock John Thomson Chen GuPaul Cliften (Geno Schupp) Mike Barker Matt Kusner
Other supportive roles (e.g. research or editorial assistance, samples, scripts, data, photographs, etc.)Other supportive roles (e.g. research or editorial assistance, samples, scripts, data, photographs, etc.)
Norm Wickett Paul OvervoordeOther supportive roles (e.g. research or editorial assistance, samples, scripts, data, photographs, etc.)Other supportive roles (e.g. research or editorial assistance, samples, scripts, data, photographs, etc.) Funding Agencies:Funding Agencies:
Katrina Dlugosch Ninglin Yin Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology Center
Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology Center
Eric Wafula Jacob Parnell
Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology Center
Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology CenterJacob Davidson Keithanne Mockaitis
Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology Center
Utah State University: Department of Biology; Vice President for Research; Center for Integrated Biosystems; Center for High Performance Computing; Ecology Center
Alan Smith Elizabeth Sheffield National Science FoundationNational Science Foundation
Jeffrey Boore Tom Ranker Genomics Education Partnership, Howard Hughes Medical Institute and The Genome Center at Washington University
Genomics Education Partnership, Howard Hughes Medical Institute and The Genome Center at Washington UniversityBrent Mishler Laura Forrest
Genomics Education Partnership, Howard Hughes Medical Institute and The Genome Center at Washington University
Genomics Education Partnership, Howard Hughes Medical Institute and The Genome Center at Washington University
Kenneth White Brian Joy Dedication: Kristal, Cora, and TylerThis one’s for you. Thanks.Dedication: Kristal, Cora, and TylerThis one’s for you. Thanks.Daryll DewaldDedication: Kristal, Cora, and TylerThis one’s for you. Thanks.Dedication: Kristal, Cora, and TylerThis one’s for you. Thanks.
53Friday, May 28, 2010
Photo Credits: fruit fly: André Karwath; mosquito: Ashok Prabhakaran; honeybee: Allie Caulfield; sea urchin: Jerry Kirkhart; green anole lizard: Clinton & Charles Robertson; opossum: Mike Keeling; platypus: Jon Hooper; horse: Pete Birkinshaw; cow: Kevek Law; rhesus macaque: Paul Asman & Jill Lenoble; nematode: Flickr user snickclunk; dog: John Wright; sea squirt: Silke Baron; chicken: Ernst Vikne; human: Lindsey Wilson; chimp: Frank Wouters; mouse: Kim Carpenter; pufferfish: Julien Mouille; Arabidopsis: Sui-setz; rice: FotoosVanRobin; grape: Tomomarusan; papaya: geishaboy500; yeast: David O. Morgan; diatom: Minami Himemiya; giant panda: Aaron Logan; black cottonwood: Dave Powell; woolly mammoth: Mauricio Anton; wine glass: Klaus Post
Collage by Kristal Watrous
54Friday, May 28, 2010