Actinophryid


The actinophryids are an order of heliozoa, a polyphyletic array of stramenopiles, having a close relationship with pedinellids and Ciliophrys. They are common in fresh water and occasionally found in marine and soil habitats. Actinophryids are unicellular and roughly spherical in shape, with many axopodia that radiate outward from the cell body. Axopodia are a type of pseudopodia that are supported by hundreds of microtubules arranged in interlocking spirals and forming a needle-like internal structure or axoneme. Small granules, extrusomes, that lie under the membrane of the body and axopodia capture flagellates, ciliates and small metazoa that make contact with the arms.[1][2]

Actinophryids are largely aquatic protozoa with a spherical cell body and many needle-like axopodia. They resemble the shape of a sun due to this structure, which is the inspiration for their common name: heliozoa, or "sun-animalcules". Their bodies, without arms, range in size from a few tens of micrometers to slightly under a millimeter across.[3]

The outer region of cell body is often vacuolated. The endoplasm of actinophryids is less vacuolated than the outer layer, and a sharp boundary layer may be seen by light microscopy.[4] The organisms can be either mononucleate, with a single, well defined nucleus in the center of the cell body, or multinucleate, with 10 or more nuclei located under the outer vacuolated layer of cytoplasm. The cytoplasm of actinophryids is often granular, similar to that of Amoeba.[5]

Actinoprhyid cells may fuse when feeding, creating larger aggregated organisms. Fine granules that occur just under the cell membrane are used up when food vacuoles form to enclose prey.[6] Actinophryids may also form cysts when food is not readily available. A layer of siliceous plates is deposited under the cell membrane during the encystment process.[7]

Contractile vacuoles are common in these organisms, which are presumed to use them to maintain body volume by expelling fluids to compensate for the entry of water by osmosis. Contractile vacuoles are visible as clear bulges from the surface of the cell body that slowly fill then rapidly deflate, expelling their contents into the environment.

The most distinctive characteristic of the actinophryids is their axopodia. These axopodia consist of a central, rigid rod which is coated in a thin layer of ectoplasm. In Actinophrys the axonemes end on the surface of the central nucleus, and in the multicellular Actinosphaerium they end at or near nuclei.[5] The axonemes are composed microtubules arranged in a double spiral pattern characteristic of the order.[8] Due to their long, parallel construction these microtubules demonstrate strong birefringence.[9][10]


Video of a contractile vacuole collapse in Actinosphaerium
Cross-section of the double spiral microtubule structure in an axopod
Actinophrys undergoing multiple plasmotomy