Biogenesis of peroxisomes and mitochondria linked by division

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Histochem Cell Biol, function of the ER in the indirect delivery of most or all directly transferred from the ER to peroxisomes by a. PMPs Tabak et al 2008 to an ER function in comprising nonvesicular pathway suggesting that ER to peroxisome. some few but essential PMPs e g Pex3p and Pex16p vesicular transport is not required to provide lipids for. semi autonomous model Haan et al 2006 Hoepfner peroxisomal growth Raychaudhuri and Prinz 2008 It is. et al 2005 Kim et al 2006 Kragt et al 2005 Motley and long known from ultrastructural studies that peroxisomes. Hettema 2007 Mullen and Trelease 2006 Tam et al 2005 are often found in close contact with the ER Thus speciWc. The ER maturation model received strong support by the ER subdomains contact sites might mediate the transport. observation that a loss of the peroxins Pex3p Pex16p or of phospholipids to the growing peroxisomes by a transfer. Pex19p which are required for peroxisomal membrane bio mechanism that has yet to be elucidated. genesis and PMP targeting insertion Van Ael and Fransen. 2006 resulted in the absence of detectable peroxisomes. peroxisomal membranes whereas reintroduction led to a de Division of peroxisomes. novo formation of peroxisomes from the ER The Pex3p. and Pex19p have been observed to initially localize to the In the last few years great progress has been made in the. ER before maturing into import competent peroxisomes identiWcation of factors involved in the division of peroxi. Haan et al 2006 Hoepfner et al 2005 Tam et al 2005 somes Table 1 Fig 1 First dynamin like proteins. indicating that the ER is the source of the newly synthe DLPs namely yeast Vps1 and mammalian DLP1 Drp1. sized membrane and organelle However recent Wndings in were shown to be involved in the division step of peroxi. S cerevisiae indicate that in wild type cells peroxisomes somal biogenesis Hoepfner et al 2001 Koch et al 2003. multiply by growth and division and do not form de novo Li and Gould 2003 Dynamins are large GTPases known. Only cells lacking peroxisomes as a result of a segregation to perform Wssion of tubulated membranes Praefcke and. defect were observed to form peroxisomes de novo out of McMahon 2004 In line with this silencing of DLP1 leads. the ER Motley and Hettema 2007 Nagotu et al 2008b to highly elongated peroxisomal tubules which are con. Furthermore evidence has been presented that Pex3p is stricted but cannot be divided Koch et al 2004 Interest. directly transported to peroxisomes in a novel Pex19p and ingly DLP1 is also involved in mitochondrial division. Pex16p dependent manner in mammalian cells Matsuzaki reviewed in Hoppins et al 2007 Okamoto and Shaw. and Fujiki 2008 Thus the physiological signiWcance of 2005 and was therefore the Wrst component identiWed to. the mechanism of de novo formation in comparison to the divide two diVerent organelles Fig 1 Furthermore mam. classical pathway of growth and division is debatable The malian hFis1 Fission 1 a tail anchored protein of the. de novo formation of peroxisomes from the ER may repre outer mitochondrial membrane reviewed in Hoppins et al. sent a rescue mechanism that becomes functional in some 2007 Okamoto and Shaw 2005 was likewise found on. organisms only in case peroxisomes are lost e g due to peroxisomes and its involvement in peroxisomal division. failure in inheritance Most of the studies addressing de was demonstrated Kobayashi et al 2007 Koch et al 2005. novo peroxisome formation out of the ER have been per Table 1 Fig 1 The hFis1 is supposed to act as an adaptor. formed in yeast fungi and at present there is only one molecule for the recruitment of DLP1 and regulate orga. report in mammalian cells Kim et al 2006 It is interest nelle morphology and Wssion through self interaction Ser. ing to note that in yeast PMPs are often misdirected to the asinghe and Yoon 2008 An increase of hFis1 promotes. ER under conditions of peroxisomal dysfunction whereas mitochondrial and peroxisomal division while its loss of. in mammalian cells they are mistargeted to mitochondria function inhibits the division of both organelles Kobayashi. Van Ael and Fransen 2006 The concept of an indirect tar et al 2007 Koch et al 2005 Based on these Wndings the. geting of PMPs from the ER to peroxisomes has brought up existence of other shared components of peroxisomal and. several yet unanswered questions This novel traYcking mitochondrial division has been proposed Schrader 2006. pathway is supposed to involve ER derived vesicular or Very recently the mammalian tail anchored protein mito. pre peroxisomal structures which either fuse with existing chondrial Wssion factor MV was identiWed to be the third. peroxisomes or develop into mature organelles Motley and factor involved in mitochondrial and peroxisomal division. Hettema 2007 Tabak et al 2008 Current studies in the Gandre Babbe and van der Bliek 2008 Table 1 Fig 1. Weld try to elucidate if and how the PMPs enter and leave Similar to hFis1 loss of MV function results in the elonga. the ER how they are sorted and packaged what the nature tion of both organelles The MV is supposed to act as an. and composition of the ER derived structures is how adaptor protein as well but in a complex diVerent from. fusion with peroxisomes is mediated and if a retrograde hFis1 It is possible that MV is involved in the assembly of. peroxisome to ER pathway exists Halbach et al 2008 the constriction machinery Components involved in orga. Matsuzaki and Fujiki 2008 Mullen and Trelease 2006 In nelle constriction prior to Wssion have so far not been iden. a recent report it has been demonstrated that lipids are tiWed and the process is currently not well understood. Histochem Cell Biol,Table 1 Shared components of,Plants Yeast Mammals. the peroxisomal and mitochon, drial division machinery across Peroxisomes Mitochondria Peroxisomes Mitochondria Peroxisomes Mitochondria Family Function. organisms Membrane,TA protein adapter, Fis1a b Fis1a b Fis1 Fis1 hFis1 hFis1 TPR motif protein. Mff Mff TA protein adapter,a a Mdv1 Mdv1 WD protein.
IdentiWed in A thaliana ELM,Arimura et al 2008 protein. b DRP3A B DRP3A B Dnm1 Dnm1 Drp1 DLP1 Drp1 DLP1 large Final. Only present in S cerevisae c,DRP1C E Vps1 GTPase scission. and C glabrata,c Mmm1 2 Membrane,Required in S cerevisae but Pex11 Pex11. Pex11 Mdm 10 12 tubulation,not in H polymorpha 31 33. Fig 1 Organelle division in mammals Peroxisomes and mitochon abundance and to tubulate membranes MV is a novel TA protein. dria are divided by similar machineries DLP1 is a GTPase performing involved in peroxisomal and mitochondrial division It is supposed to. the Wnal scission of constricted membranes and is recruited from the function in a complex diVerent from hFis1 In yeast additional linker. cytosol by the TA protein hFis1 which might regulate organelle mor proteins Mdv1 Caf4 have been identiWed which are involved in mito. phology and Wssion through self interaction hFis1 is supposed to inter chondrial and peroxisomal division see Table 1. act with the peroxin Pex11 which is known to regulate peroxisomal. Schrader and Fahimi 2006 Alternatively hFis1 might identiWed The major DLP required for peroxisomal Wssion. play a role in membrane constriction Serasinghe and Yoon in glucose grown S cerevisiae appears to be vacuolar pro. 2008 tein sorting associated protein 1 Vps1 Hoepfner et al. Sharing the key components of the division machinery 2001 The Vps1 was initially shown to be required for vac. appears to be a ubiquitous and general principle of peroxi uolar protein sorting but is not involved in mitochondrial. somes and mitochondria in many eukaryotes as homolo Wssion However cells grown in peroxisome inducing. gous components were identiWed to act on both organelles growth conditions oleate have been reported to require the. in fungi and plants Table 1 Members of the DRP3 sub yeast DLP Dnm1 yeast Fis1 and the soluble molecular. family of DLPs as well as the Fis1 homologs Fis1a and linkers Mdv1 and the closely related Caf4 Kuravi et al. Fis1b have been implicated in peroxisomal and mitochon 2006 Motley and Hettema 2007 Motley et al 2008. drial division in Arabidopsis thaliana Lingard et al 2008 Table 1 Mdv1 and Caf4 are cytosolic WD proteins that. Mano et al 2004 Zhang and Hu 2008 In yeast two inde bind to Fis1 and Dnm1 in yeast In addition the four pro. pendent pathways of peroxisomal division have been teins are key components of mitochondrial division in yeast. Histochem Cell Biol, reviewed in Hoppins et al 2007 Okamoto and Shaw peroxisomes or mitochondria and how is their division.
2005 Dnm1 but not Vps1 Fis1 and Mdv1 are also coordinated How are membrane proteins such as Fis1 and. required for peroxisomal and mitochondrial Wssion in MV targeted to both peroxisomes and mitochondria It is. H polymorpha Nagotu et al 2008a Nagotu et al 2008b likely that there is already regulation at the targeting level. Interestingly Caf4 proteins appear to be present only in of these proteins to both organelles As hFis1 recruits DLP1. Saccharomyces species and Candida glabrata but not in to mitochondria and peroxisomes Yu et al 2005 the. other yeast and fungi including H polymorpha and hFis1 levels in the respective membrane might be critical. P pastoris However all species tested contain Dnm1 for the regulation of division Recently it was shown that. Vps1 as well as Fis1 orthologs Nagotu et al 2008a The the targeting of hFis1 to peroxisomes depends on Pex19p. de novo formation of peroxisomes via the ER see above an import factor involved in peroxisomal membrane protein. appears to be independent of Vps1 and Dnm1 Motley and import Delille and Schrader 2008 In contrast targeting. Hettema 2007 Nagotu et al 2008b of hFis1 to mitochondria does not require any known. Interestingly there are no homologs for the linker pro mitochondrial transport components Kemper et al 2008. teins Mdv1 or Caf4 in metazoans while MV appears to be Setoguchi et al 2006 and appears to be independent from. metazoan speciWc Gandre Babbe and van der Bliek 2008 peroxisomal targeting Furthermore DLP1 is subject to. Table 1 It is an interesting question if other Fis1 interact several posttranslational modiWcations such as phosphory. ing linker proteins functionally similar to Mdv1 and Caf4 lation ubiquitination and sumoylation which contribute to. exist in higher eukaryotes which might be involved in the its regulation reviewed in Cerveny et al 2007. speciWc regulation and assembly of the division machiner. ies on mitochondria and peroxisomes, Besides the overlap in some components of the peroxi Interrelationship of peroxisomes and mitochondria. somal and mitochondrial division machinery there is so far. no evidence for shared components in other steps of their There is emerging new evidence that peroxisomes and. biogenesis Membrane proteins of the Pex11 family have mitochondria exhibit a far greater co dependent relation. been shown to induce proliferation of peroxisomes by elon ship than previously assumed see also Schrader and Yoon. gation growth of the organelle membrane and subsequent 2007 Peroxisomes and mitochondria are metabolically. division in mammals yeast and plants Lingard et al 2008 linked organelles They cooperate in the oxidation of. Schrader and Fahimi 2006 Thoms and Erdmann 2005 Yan fatty acids and in the metabolism of reactive oxygen spe. et al 2005 Table 1 Fig 1 However the mode of action cies In addition a novel vesicular transport pathway from. of the Pex11 proteins is still not fully understood In mam mitochondria to peroxisomes has been described Neuspiel. mals only three Pex11 proteins and have been et al 2008 Therefore it is meaningful that their biogene. reported to be involved in the regulation of peroxisome sis is coordinated and that they share components of their. abundance proliferation whereas in yeast several peroxins division machinery Schrader 2006 Table 1 Fig 1. aVecting peroxisome abundance have been identiWed It has Moreover peroxisomal alterations in metabolism biogene. been shown that the Pex11 proteins interact with the sis dynamics and proliferation can potentially inXuence. respective Fis1 proteins but not with the DLPs Kobayashi mitochondrial functions and vice versa The close interrela. et al 2007 Koch et al 2005 Lingard et al 2008 The tionship between peroxisomes and mitochondria is not only. function of this interaction is still unclear Interestingly supposed to have an impact on their cooperative functional. several mitochondrial membrane proteins required for the ity but might also contribute to diseases Camoes et al. formation of mitochondrial tubules have been identiWed in 2008 Peroxisomal and mitochondrial dysfunctions are. fungi but not yet in higher eukaryotes reviewed in Hoppins often associated with neurological and developmental. et al 2007 Okamoto and Shaw 2005 Table 1 defects Furthermore peroxisomes and mitochondria have. There is great interest now in the identiWcation of further been suggested to contribute to pathological conditions. components involved in peroxisomal and or mitochondrial associated with oxidative stress and have been linked to. division It is likely that the dynamic multistep process of ageing as well Terlecky et al 2006 In line with this it has. peroxisome elongation growth constriction Wnal scis recently been shown that elevating peroxisomal catalase. sion and proper intracellular distribution via the cytoskele levels in late passage human cells restores mitochondrial. ton and motor proteins involves the assembly of distinct integrity Koepke et al 2007 whereas compromised cata. Histochem Cell Biol DOI 10 1007 s00418 009 0561 9 123 REVIEW Biogenesis of peroxisomes and mitochondria linked by division Hannah K Delille Renato Alves Michael Schrader

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