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==Characteristics== Like other [[diplomonad]]s, ''Giardia'' have two [[cell nucleus|nuclei]], each with four associated [[Flagellum|flagella]], and were thought to lack both [[mitochondrion|mitochondria]] and a [[Golgi apparatus]]. However they are now known to possess a complex endomembrane system as well as mitochondrial remnants, called [[mitosome]]s, through mitochondrial reduction.<ref>{{Cite journal|last1=Tovar|first1=Jorge|last2=León-Avila|first2=Gloria|last3=Sánchez|first3=Lidya|last4=Sutak|first4=Robert|last5=Tachezy|first5=Jan|last6=van der Giezen|first6=Mark|last7=Hernández|first7=Manuel|last8=Müller|first8=Miklós|last9=Lucocq|first9=John|date=2003|title=Mitochondrial remnant organelles of Giardia function in iron-sulphur protein maturation|journal=Nature|volume=426|issue=6963|pages=172–176|doi=10.1038/nature01945|pmid=14614504|bibcode=2003Natur.426..172T|s2cid=4402808}}</ref> <ref>{{cite journal | author= Anna Karnkowska| title= A Eukaryote without a Mitochondrial Organelle | journal= Current Biology| date= May 2016| url=http://www.cell.com/current-biology/fulltext/S0960-9822(16)30263-9|display-authors=etal |doi=10.1016/j.cub.2016.03.053 | pmid= 27185558 |volume=26 | issue= 10 |pages=1274–1284| doi-access= free }} </ref> <ref>{{cite journal |vauthors=Soltys BJ, Falah M, Gupta RS |title=Identification of endoplasmic reticulum in the primitive eukaryote ''Giardia lamblia'' using cryoelectron microscopy and antibody to Bip |journal=J. Cell Sci. |volume=109 |issue=Pt 7 |pages=1909–17 |date=July 1996 |doi=10.1242/jcs.109.7.1909 |pmid=8832413 |url=http://jcs.biologists.org/cgi/pmidlookup?view=long&pmid=8832413}}</ref><ref>{{cite journal |author1=Dolezal P |author2=Smíd O |author3=Rada P |display-authors=etal |title=Giardia mitosomes and trichomonad hydrogenosomes share a common mode of protein targeting |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=102 |issue=31 |pages=10924–9 |date=August 2005 |pmid=16040811 |pmc=1182405 |doi=10.1073/pnas.0500349102 |bibcode=2005PNAS..10210924D }}</ref> The mitosomes are not used in [[Adenosine triphosphate|ATP]] synthesis the way mitochondria are, but are involved in the maturation of [[iron-sulfur protein]]s.<ref name="Tovar2003">{{cite journal |vauthors=Tovar J, etal | year = 2003 | title = Mitochondrial remnant organelles of Giardia function in iron-sulphur protein maturation | journal = Nature | volume = 426 | issue = 6963 | pages = 172–6 | pmid = 14614504 | doi = 10.1038/nature01945 | bibcode = 2003Natur.426..172T | s2cid = 4402808 }}</ref> The [[synapomorphy|synapomorphies]] of genus ''Giardia'' include cells with duplicate organelles, absence of [[cytostome]]s, and ventral adhesive disc.<ref>{{cite web |last=Cepicka |first=Ivan |title=Fornicata |website=Tree of Life Web Project |date=September 2008 |url=http://tolweb.org/Fornicata/121182/2008.09.02}}</ref>
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