Algal Intrafamilial Strife

Most of you are probably familiar with the old adage that one should keep one's friends close and one's enemies closer. From a phylogenetic perspective, the red algal genus Plocamium has certainly achieved the latter.

Plocamium species growing on the coast of South Africa, copyright Derek Keats.

Plocamiaceae is a cosmopolitan family of marine red algae found mostly in temperate waters. They may grow in a variety of habitats from sheltered to exposed. Phylogenetic analyses have indicated that the family is somewhat distantly related to other red algal families, such that it is currently classified in its own order (Saunders & Kraft 1994). The great majority of the forty-odd known species of Plocamiaceae are currently placed in the genus Plocamium. These are reasonably sized seaweeds with erect or decumbent thalli that can grow about half a metre in length/height. They have flattened, complanately branched axes (that is, the branches are in the same plane as the axis they branch from). Branching is pectinate (comb-like) with each axis producing usually between two and six branchlets. The lower branchlets in a series are usually unbranched but higher ones will produce their own series of side-branchlets. In particular, the last branchlet will generally grow and overtop the axis it arose from to effectively replace it (as a result, the axis of the algal thallus will appear at first glance to have many more side branches than mentioned previously but can be seen on close inspection to have something of a zig-zag appearance representing the successive axes). The comb-like pattern of the branching is particularly evident in terminal branches of the thallus. In section, the axes have a disorganised cortex surrounding the central axial cells. Plocamiaceae have the standard triphasic red algal life cycle with gametophytes and sporophytes similar in outward appearance. Cystocarps appear to be more or less globular and borne along axial margins. Tetrasporangia are borne on the underside of modified branchlets called stichidia in a manner reminiscent of the sporangia of ferns (Gabrielson & Scagel 1989).

Close-up on terminal brachlets of Plocamium coccineum, copyright Fernan Federici. Some tetrasporangia-bearing stichidia are visible in the lower part of the image.

Only a few species have been described to date of the other genus of Plocamiaceae, Plocamiocolax. Though its reproductive anatomy demonstrates its relationship to Plocamium, Plocamiocolax is very different in its superficial appearance. It is a parasite, specifically a parasite of its sister genus. As such, they exhibit greatly reduced thalli and coloration. They grow on the host Plocamium as wartlike cushions, up to about five millimetres in diameter. As the cushion grows, it produces short, flattened projections that may be simple or forked. Tetrasporophytes may bear tetrasporangia on greatly reduced stichidia or on partially endophytic, verrucose patches.

Plocamiocolax pulvinata growing on Plocamium, copyright Michael Hawkes.

Parasitic forms that are closely related to the species they infect are referred to as 'adelphoparasites', meaning 'sister-parasites'. Adelphoparasitism is remarkably common among red algae: of over sixty known genera of parasitic red algae, about 90% are adelphoparasites (Salomaki & Lane 2014). One of the very first posts I ever wrote on this site was about red algal adelphoparasites, way back in...(gosh, really?...doesn't time fly when you're marching unceasingly towards oblivion...) It is possible that the regularity of this phenomenon is related to a distinctive feature of red algal development: the ability to open connections between adjacent cells allowing the passage of cytoplasm and organelles. Though the primary function of this process is presumably to facilitate the transfer of cellular products between cells of a single individual, it is not difficult to imagine a scenario where one individual hijacks another. The more closely related the adjacent cells, the greater the chance of an illicit connection succeeding. And succeeding multiple times. Though treated as a distinct genus, 'Plocamiocolax' lineages have apparently arisen within Plocamium multiple times (Goff et al. 1996). In some cases, a Plocamiocolax species proves to be the direct derivative of the Plocamium species they are found infesting. In others, a Plocamiocolax has arisen on one host species but later made the switch to another. Children are supposed to become independent and find their own way in the world, but sometimes the blighters just won't leave.

REFERENCES

Gabrielson, P. W., & R. F. Scagel. 1989. The marine algae of British Columbia, northern Washington, and southeast Alaska: division Rhodophyta (red algae), class Rhodophyceae, order Gigartinales, families Caulacanthaceae and Plocamiaceae. Canadian Journal of Botany 67: 1221–1234.

Goff, L. J., D. A. Moon, P. Nyvall, B. Stache, K. Mangin & G. Zuccarello. 1996. The evolution of parasitism in the red algae: molecular comparisons of adelphoparasites and their hosts. Journal of Phycology 32: 297–312.

Salomaki, E. D., & C. E. Lane. 2014. Are all red algal parasites cut from the same cloth? Acta Societatis Botanicorum Poloniae 83 (4): 369–375.

Saunders, G. W., & G. T. Kraft. 1994. Small-subunit rRNA gene sequences from representatives of selected families of the Gigartinales and Rhodymeniales (Rhodophyta). 1. Evidence for the Plocamiales ord.nov. Canadian Journal of Botany 72: 1250–1263.