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Further studies have also revealed unique stage-specific roles of these kinases, but fully understanding several aspects of their biological function has been thwarted by their essential functions and by the fact that Apicomplexa are obligate intracellular parasites that cannot be cultivated axenically. In coccidian parasites, including T.

Initial studies suggested that the two forms were functionally redundant 5 , but these studies were performed using genetic techniques that relied upon overexpression of a second copy of the gene; at the time, precise conditional expression of PKG was not technically feasible.

Since the original studies investigating PKG biology were reported, the repertoire of tools available for manipulation of gene expression in the Apicomplexa has expanded dramatically. Clustered regularly interspaced short palindromic repeat CRISPR -Cas9 gene-editing technology has revolutionized the field and enabled rapid generation of precise mutations, and now it is possible to create mutants that express a target protein in the correct genomic context and create transgenic parasites in nearly any genetic background.

For essential genes, conditional expression is required to understand the molecular function of genes.

Both tetracycline-regulated transcriptional systems and posttranslational conditional expression systems using the FKbinding protein FKBP -inducible degradation domain DD have been developed for T. In addition, because the DD-stabilizing agent Shield must be continuously applied to stabilize proteins, the DD system can be expensive and cumbersome.

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The auxin-inducible degron AID system has improved kinetics and responsiveness in T. The F box recruits specific substrates for degradation by the proteosomal system. Auxin stabilizes the interaction of the auxin receptor TIR1 an F box protein with proteins that encode an auxin-inducible degron AID motif 7. To use AID, Brown et al. Pretreatment with proteasome inhibitors resulted in stabilization of the YFP-mAID, confirming the role of the proteasome in protein depletion.

Auxin had no effect on parasite replication, suggesting a role for PKG in motility, egress, or invasion. Complementation of PKG-mAID-3HA parasites with different PKG mutants revealed that the membrane-associated form was sufficient for critical functions, including microneme secretion and invasion as well as parasite egress. Parasite lines complemented with cytosolic forms of PKG were unable to make plaques and had reduced capacity for microneme secretion, invasion, and egress.

The PKG cytosolic form was dispensable for parasite growth, indicating that, in tachyzoites, only the membrane-associated form of PKG is critical for parasite viability 6.

Ubiquitination is common among T. The utility of the AID system provides functional evidence that the mechanics of the SCF ubiquitin ligase machinery are conserved in T. A number of T. As reported for PKG, acylation usually requires N-terminal myristoylation followed by palmitoylation.

Since acylation and proper targeting are critical for the essential functions of PKG, development of compounds that target membrane trafficking of PKG may pose an alternative strategy for development of antiparasitic agents. Protein acylation also is common in the Kinetoplastida 9 , and thus acylation enzymes N-myristoyl transferases or palmitoyl transferases may prove to be potential general targets for treatment of parasitic infections.

In the tissue-cyst forming coccidian Apicomplexa T. By analogy, a role for cytosolic forms of T.

The TIR1 stable strain was made in the RH background, which is unable to differentiate to bradyzoites or to complete the full life cycle in feline intestines. Assuming that TIR1 strains are readily made in other strains of T. In addition, application of AID may enable dissection of the molecular mechanisms of complex cell signaling pathways in T.

Philip and Waters administered auxin to synchronized, in vitro-cultivated, PbCnA-AID parasites at specific points in Plasmodium development and performed a battery of functional and morphological assays These assays demonstrated a role for calcineurin in erythrocyte attachment and invasion, male gametocytogenesis, and fertilization, as well as ookinete-to-oocyst and sporozoite-to-liver-stage transitions Neither Cryptosporidium nor T.

AID, adapted for use in mouse or organoid models, may also be a useful tool for regulated expression of genes in apicomplexan species or life cycle stages that were formerly inaccessible to experimental manipulation. Notes The views expressed in this Commentary do not necessarily reflect the views of this journal or of ASM.

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