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Summary  Egress of malaria parasites from the host cell requires the concerted rupture of its enveloping membranes. Hence, we investigated the role of the plasmodial perforin‐like protein  PPLP 2 in the egress of   P   lasmodium falciparum  from erythrocytes.  PPLP 2 is expressed in blood stage schizonts and mature gametocytes. The protein localizes in vesicular structures, which in activated gametocytes discharge  PPLP 2 in a calcium‐dependent manner.  PPLP 2 comprises a  MACPF  domain and recombinant  PPLP 2 has haemolytic activities towards erythrocytes.  PPLP 2‐deficient [ PPLP 2(−)] merozoites show normal egress dynamics during the erythrocytic replication cycle, but activated  PPLP 2(−) gametocytes were unable to leave erythrocytes and stayed trapped within these cells. While the parasitophorous vacuole membrane ruptured normally, the activated  PPLP 2(−) gametocytes were unable to permeabilize the erythrocyte membrane and to release the erythrocyte cytoplasm. In consequence, transmission of  PPLP 2(−) parasites to the  Anopheles  vector was reduced. Pore‐forming equinatoxin  II  rescued both  PPLP 2(−) gametocyte exflagellation and parasite transmission. The pore sealant Tetronic 90R4, on the other hand, caused trapping of activated wild‐type gametocytes within the enveloping erythrocytes, thus mimicking the  PPLP 2(−) loss‐of‐function phenotype. We propose that the haemolytic activity of  PPLP 2 is essential for gametocyte egress due to permeabilization of the erythrocyte membrane and depletion of the erythrocyte cytoplasm.

Perforin‐like protein PPLP 2 permeabilizes the red blood cell membrane during egress of P lasmodium falciparum gametocytes