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Most eukaryotic parasites are obligately heteroxenous, requiring sequential infection of different host species in order to survive.  Toxoplasma gondii  is a rare exception to this rule, having a uniquely facultative heteroxenous life cycle. To understand the origins of this phenomenon, we compared development and stress responses in  T. gondii  to those of its its obligately heteroxenous relative,  Hammondia hammondi  and have identified multiple  H. hammondi  growth states that are distinct from those in  T. gondii.  Of these, the most dramatic difference was that  H. hammondi  was refractory to stressors that robustly induce cyst formation in  T. gondii,  and this was reflected most dramatically in its unchanging transcriptome after stress exposure. We also found that  H. hammondi  could be propagated  in vitro  for up to 8 days post-excystation, and we exploited this to generate the first ever transgenic  H. hammondi  line. Overall our data show that  H. hammondi  zoites grow as stringently regulated, unique life stages that are distinct from  T. gondii  tachyzoites, and implicate stress sensitivity as a potential developmental innovation that increased the flexibility of the  T. gondii  life cycle.

elife

Dissection of the  in vitro  developmental program of  Hammondia hammondi  reveals a link between stress sensitivity and life cycle flexibility in  Toxoplasma gondii