Back to Moss Dub for this post because Chris Carter has sent me some stunning images of the filamentous desmid Desmidium grevillei that I talked about in my earlier post. I mentioned that it is surrounded by a mucilaginous sheath, which was just apparent in my brightfield image. Chris has added Indian ink to the wet mount. The ink forms a dense suspension in the water but is repelled by the mucilage around the desmid cells, resulting in a much better impression of the extent of the sheath around the cell than is otherwise possible.
Desmidium grevillei from Moss Dub, photographed by Chris Carter using Indian ink to highlight the mucilage sheath around the cells.
Indian ink is a negative stain, which means that it is the background, rather than the specimen itself, which takes up the colour. This, in turn, alters the passage of light through the sample and appears to improve the contrast of the final image. Chris’ images of the apical view show this well, and also illustrates the complicated three-dimensional arrangement of the chloroplasts within each semi-cell. His photographs also show the pores through which the mucilage is secreted.
The curious thing about this negative stain is that, whilst it appears to emphasis a halo of nothingness around the Desmidium filament, it is actually drawing our attention to something important. In his presidential address to the British Phycological Society in 1981 A.D. Boney referred to mucilage as “the ubiquitous algal attribute” and goes on to list the many functions that the slimes produced by a wide range of algal groups may perform. Not all will apply to our Desmidium but Boney does use desmids as examples of some of the roles slime may play: it can be, for example, a buoyancy aid, keeping the desmids in the well-lit regions of a lake or pond and it can protect cells against desiccation if a pond or lake dries out. It may also play a role in helping desmids adhere to their substrates and there is also evidence that mucilage layers may help to protect algae from toxins.
Apical view (at four different focal planes) of Desmidium grevillei from Moss Dub, photographed by Chris Carter, September 2019.
But that’s only part of the story. There is two-way traffic across the membranes of algal cells, with essential nutrients moving into the cell but, in some cases, enzymes moving in the opposite direction. If nutrients are in short supply then these enzymes can help the cell by breaking down organic molecules in order to release nutrients that can then be absorbed. Those enzymes take energy to manufacture, and the sheath of gunk around the filament means that there is a lower chance of them diffusing away before doing their job (see “Life in the colonies …”). The same principle applies to sexual reproduction too, with mucilage serving, in some cases, as “sperm traps” or simply as the phycological equivalent of KY Jelly.
It is not just the algae that benefit from this mucilage: the outer layers, especially, can be colonised by bacteria which will also be hoovering up any spare organic molecules for their own benefit with, no doubt, some collateral benefits for the organisms around them. The connection is probably too tenuous to count as a symbiosis with the desmids but we could think in terms of mutual benefits.
So that “nothing” really is a “something”, and that is before we consider the role of these extracellular compounds in the wider ecosystem. I mentioned the role of similar compounds in consolidating the fine sediments on coastal mudflats in “In the shadow of the Venerable Bede” to give a flavour of this. The least prepossessing aspect of the least prepossessing plants can, given time, change landscapes. That should give us all pause for thought.
Close-up of Desmidium grevillei filament with focus on the left-hand cell adjusted to show the apical pores. Photographed by Chris Carter from material from Moss Dub collected in September 2019.
Reference
Boney, A.D. (1981). Mucilage: the ubiquitous algal attribute. British Phycological Journal 16: 115-132.
Domozych, D. S., & Domozych, C. R. (2008). Desmids and biofilms of freshwater wetlands: Development and microarchitecture. Microbial Ecology https://doi.org/10.1007/s00248-007-9253-y
Sorentino, C. (1985). Copper resistance in Hormidium fluitans (Gay) Heering (Ulotrichaceae, Chlorophyceae). Phycologia 24: 366-388. https://doi.org/10.2216/i0031-8884-24-3-366.1