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. Author manuscript; available in PMC: 2014 Nov 17.
Published in final edited form as: Mol Reprod Dev. 2013 Jun 27;80(8):632–664. doi: 10.1002/mrd.22195

TABLE 1.

Survey of Piwi Proteins Across Animal Phylogeny: Somatic Expression Versus Germline Expression and Nuclear Expression Versus Cytoplasmic Expression

Organism Homologues Localization Expression
Germline* Soma*
Fly Piwi Cytoplasm, Nucleus PGCs
(Megosh et al, 2006)
Embryonic somatic cells
(Megosh et al, 2006)
Nucleus Ovary-GSCs, 16 cell
cysts. Oocyte, Nurse cells
(Cox et al, 1998)
Ovary - Terminal filament,
Epithelial sheath cells,
Follicle cells
(Cox et al, 1998)
Testis - GSCs, Gonialblasts
(Cox et al, 2000)
Testis - Hub cells, Somatic
stem cells, Cyst progenitor cells
(Cox et al, 2000)
Salivary Gland
(Brower-Toland et al, 2007)
Aub Cytoplasm PGCs (Harris and Macdonald, 2001) Embryonic soma (Harris and Macdonald, 2001)
Ovary - GSCs, all cyst cells,
Oocyte, Nurse cells (Brennecke et al, 2007)
Testis - GSCs, Gonialblasts,
Spermatogonia, Spermatocytes
(Nishida et al, 2007)
Ago3 Cytoplasm Ovary - Germline stem cells, All
cyst cells, Oocyte,
Nurse cells
(Gunawardane et al, 2007)
Ovary- Faint in follicle cells
(Gunawardane et al, 2007)
Cap cells
(Brennecke et al, 2007)
Testis -
GSCs, Gonialblasts, Spermatogoni
a (Nagao et al, 2010)
Mouse MIWI Cytoplasm,
Nucleus (Dense body)
(Beyret and Lin, 2011)
Testis -Meiotic spermatocytes,
Elongating spermatids
(Deng and Lin, 2002)
Unknown Pancreas (Yan et al, 2011)
Brain, Heart, Liver, Lung, Kidney
(Lee et al, 2011)
MILI Cytoplasm,
Nucleus (Dense body)
(Beyret and Lin, 2011)
Testis -
GSCs, Gonocytes, Spermatogonia,
Meiotic spermatocytes,
Spermatids (Kuramochi-Miyagawa et al. 2004, Unhavaithaya et al. 2009, Wang et al. 2009)
Unknown Mesenchymal stem cells
(Wu et al, 2010)
MIWI2 N ucleus, Cytoplasm Testis - GSCs
(Aravin et al, 2008)
Testis - Sertoli cells
(Carmell et al, 2007)
Zebrafish Ziwi Cytoplasm PGCs
(Houwing et al, 2007)
Ovary - Oogonia, Stage 1
oocytes (Houwing et al, 2007)
Testis - Spermatogonia,
Spermatocytes
(Houwing et al, 2007)
Zili Nucleus PGCs
(Houwing et al, 2008)
Embryonic soma
(Sun et al, 2010)
Cytoplasm, Nucleus Ovary - Oogonia, Stage l-IV
Oocytes (Houwing et al, 2008)
Cytoplasm Testis - Spermatogonia,
Spermatocytes, Spermatids
(Houwing et al, 2008)
Frog Xiwi Cytoplasm,
Nucleus(Mitotic/Meiotic Spindles)
Embryos Stage 1–20. Germline/Somatic separation unknown(Wilczynska et al, 2009)
Ovary - Stage I - IV occytes,
Mature oocytes
(Lau et al, 2009)
Unknown Testis (Lau et al, 2009)
Xili Unknown Embryos Stage 1–42. Germline/Somatic separation unknown
(Wilczynska et al, 2009)
Cytoplasm Ovary - Stage I - IV occytes,
Mature oocytes
(Wilczynska et al, 2009)
Unknown Testis (Wilczynska et al, 2009)
Nematode PRG1 Cytoplasm Gonad- Germline stem
cells, Mitotic/meiotic germ
cells, Mature oocytes
(Batista et al, 2008)
PRG2 Unknown Unknown Unknown
Human hiwi (piwil1) Cytoplasm(Qiao et al, 2002)
Nucleus, HEK 293T cells
(Sugimoto et al, 2007)
mRNA Testis - Spermatocytes,
Round spermatids
(Qiao et al, 2002)
Unknown mRNA Hematopoietic stem
cells (Sharma et al, 2001)
mRNA Various tissues inc.
Kidney, Heart, Brain, Liver,
Muscle, Pancreas etc.
(Sharma et al, 2001)
hili (piwiL2) Nucleus, HEK293T cells
(Sugimoto et al, 2007)
cDNA Testis
(Sasaki et al, 2003)
piwiL3 Nucleus, HEK293T cells
Sugimoto et al, 2007)
cDNA Testis
(Sasaki et al, 2003)
hiwi2 (piwiL4) Nucleus, HEK293T cells
(Sugimoto et al, 2007)
cDNA Testis
(Sasaki et al, 2003)
cDNABone marrow,
Leukocytes, Pancreas
(Sasaki et al, 2003)

mRNA Various tissues inc.
Spleen, Lung, Liver, Brain,
Heart, Kidney, Ovary etc.
(Sugimoto et al, 2007)
Sea Slug(Aplysia) Piwi Unknown Ovotestis (Rajasethupathy et al, 2012)
Nucleus Brain (Rajasethupathy et al, 2012)
Sea Urchin Seawi Cytoplasm,
Nucleus(Mitotic Spindle)
Embryonic soma
(Rodriguez et al, 2005)
Ovary - Oocytes
(Rodriguez et al, 2005)
Planarian smedwi-1 Unknown mRNA Neoblasts (Reddien et al, 2005)
smedwi- 2 Unknown mRNA Neoblasts (Reddien et al, 2005)
Polychaete Annelid
(Capitella teleta)
ct-piwi1 Unknown mRNA PGCs
(Giani et al, 2011)
mRNA Somatic cells of
embryo (Giani et al, 2011)
mRNA Larvae -
Brain, foregut, mesoderm (Giani et al, 2011)
mRNA Immature oocytes
(Giani et al, 2011)
mRNA Genital ducts
(Giani et al, 2011)
mRNA Posterior growth zone
(Giani et al, 2011)
ct-piwi2 Unknown mRNA Primordial germ cells
(Giani et al, 2011)
mRNA Embryonic soma (Giani et al, 2011)
mRNA Larvae - Brain, foregut, mesoderm (Giani et al, 2011)
mRNA Immature oocytes
(Giani et al, 2011)
mRNA Genital ducts
(Giani et al, 2011)
mRNA Posterior growth zone
(Giani et al, 2011)
Colonial
Ascidian
Piwi Nucleus, Cytoplasm Gonadal primordia of ovaries and testes
(Brown et al, 2009)
Hemocytes
(Brown et al, 2009)
Endostyle-Epithelial cells,
(Brown et al, 2009)
EfPiwiA Unknown mRNA Pluripotent stem cells - Archaeocytes
(Funayama et al, 2010)
mRNA Choanocytes
(Funayama et al, 2010)
EfPiwiB Unknown mRNA Pluripotent stem cells - Archaeocytes
(Funayama et al, 2010)
mRNA Choanocytes
(Funayama et al, 2010)
Ctenophore PpiPiwi1 Unknown mRNA Female gonad - Oocytes and nurse cells
(Alie et al, 2011)
mRNA Male gonad - Developing
spermatocytes
(Alie et al, 2011)
mRNA Somatic stem cells
of tentacle root, comb rows
and aboral sensory complex
(Alie et al, 2011)
PpiPiwi2 Unknown mRNA Female gonad - Oocytes and nurse cells
(Alie et al, 2011)
mRNA Male gonad - Developing
spermatocytes
(Alie et al, 2011)
mRNA Unidentified cells of
somatic origin
(Alie et al, 2011)
*

Analogous expression in somatic, germline tissues are placed side by side. Blank boxes indicate no experimental evidence for PIWI expression.