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Mucor circinelloides  is a zygomycete fungus and an emerging opportunistic pathogen in immunocompromised patients, especially transplant recipients and in some cases otherwise healthy individuals. We have discovered a novel example of size dimorphism linked to virulence.  M. circinelloides  is a heterothallic fungus: (+)  sex  allele encodes SexP and (−)  sex  allele SexM, both of which are HMG domain protein sex determinants.  M. circinelloides  f.  lusitanicus  ( Mcl ) (−) mating type isolates produce larger asexual sporangiospores that are more virulent in the wax moth host compared to (+) isolates that produce smaller less virulent sporangiospores. The larger sporangiospores germinate inside and lyse macrophages, whereas the smaller sporangiospores do not.  sexMΔ  mutants are sterile and still produce larger virulent sporangiospores, suggesting that either the  sex  locus is not involved in virulence/spore size or the  sexP  allele plays an inhibitory role. Phylogenetic analysis supports that at least three extant subspecies populate the  M. circinelloides  complex in nature:  Mcl ,  M. circinelloides  f.  griseocyanus , and  M. circinelloides  f.  circinelloides  ( Mcc ).  Mcc  was found to be more prevalent among clinical  Mucor  isolates, and more virulent than  Mcl  in a diabetic murine model in contrast to the wax moth host. The  M. circinelloides sex  locus encodes an HMG domain protein (SexP for plus and SexM for minus mating types) flanked by genes encoding triose phosphate transporter (TPT) and RNA helicase homologs. The borders of the  sex  locus between the three subspecies differ: the  Mcg sex  locus includes the promoters of both the TPT and the RNA helicase genes, whereas the  Mcl  and  Mcc sex  locus includes only the TPT gene promoter. Mating between subspecies was restricted compared to mating within subspecies. These findings demonstrate that spore size dimorphism is linked to virulence of  M. circinelloides  species and that plasticity of the  sex  locus and adaptations in pathogenicity have occurred during speciation of the  M. circinelloides  complex.

Sporangiospore Size Dimorphism Is Linked to Virulence of Mucor circinelloides