Spheres of influence

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.


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


Desmids from Moss Dub


I’d like to say that this post is about an excursion I made beyond Ennerdale Water and along the valley of the River Liza in order to find some different habitats from those that I usually write about in this blog.  I’d like to but, in truth, I was sitting by a road about ten miles away waiting for roadside assistance whilst my compatriots on the Quekett Microscopy Club / British Phycological Society algae weekend went up the valley on a glorious mid-September afternoon without me.  Whilst I was sitting waiting for a tyre to be replaced in a garage in Egremont they were casting plankton nets and squeezing handfuls of Sphagnum beside Moss Dub, a small tarn set amidst woodland close by the River Liza.

Moss Dub is set within one of Britain’s oldest and most ambitious rewilding schemes, Wild Ennerdale, where nature is allowed to shape the landscape as far as possible free from human interference.  However, Moss Dub, as we found out, is far from a natural water body.  A path forks and the two arms act as bunds encompassing a shallow pond, now partly overgrown with aquatic vegetation.   There is evidence of past mining activity – for iron and copper – in the area and my guess is that Moss Dub was, in the far past, a reservoir associated with the Lingmell mine located on the hillside above the River Liza and active in the late 19thcentury.     Whatever its history, it proved to be a rich location for desmids, and we spent a happy Saturday dipping Pasteur pipettes into the vials of peaty water that they collected and peering through our microscopes (If you want to know more about how to collect desmids, look at the post I wrote after our last excursion to the Lake District: “Desmid masterclass”).

There were some conspicuous green growths suspended in the water at the margin of the pond.  Even without a microscope, their filamentous nature was obvious.  When magnified, we saw chains of green cells set within a distinct mucilaginous sheath.  Each filament was composed of short cells with a distinct notch on either side.   This is a representative of Desmidium, one of a relatively small number of filamentous desmids.  We met D. schwartziion our previous excursion (see “Lessons from School Knott Tarn”); that species was present here along with D. grevillei, which is similar in many respects but the cross-section is lemon-shaped rather than triangular.


Desmidium grevillei from Moss Dub, Ennerdale Valley, September 2019.  a. shows a macroscopic view of filaments in a Petri dish; b. shows a filament of cells, along with a distinct mucilaginous sheath whilst c. shows a cell in cross-section.  I forgot to bring my graticule so cannot add scale bars to any of the images in this post.   Instead, I will quote dimensions from the Freshwater Algal Flora of Britain and Ireland to give an indication of size.   Cells of D. grevillei are 30 – 56 micrometres wide (50 micrometres is 1/20thof a millimetre).   The photo at the top of this post is a view of Moss Dub, taken by Zeneb Henderson

There were numerous other desmids in the sample.  A couple are illustrated below, and we’ve sent the sample off to David Williamson for a more thorough examination, and some definitive names.  On the right-hand side of the plate there is a different green alga, Coelastrum pulchrum, a member of the Chlorophyceae that forms spherical colonies with a fixed number of cells (“coenobia”).  We met Coelastrum microporum in the River Wear last summer (see “More green algae from the River Wear”): cells of C. pulchrum, by contrast, have a blunt projection.


More algae from Moss Dub: d. Micrasterias radiosa (142 – 191 micrometres across); e. Euastrum pinnatum (65-75 micrometres across; 125 – 170 long); f. Coelastrum pulchrum (about 100 micrometres in diameter). 

The final desmid I’ve illustrated is Closterium lunula, large by desmid standards as it can reach half a millimetre or more in length.  Members of this genus have prominent vacuoles at each end of the cell within which small crystals can be seen.  Because C. lunula is so large it is easy to see both vacuole and watch Brownian motion move the crystals within.   Studies have shown that these are crystals of barium sulphate and also that the crystals are scattered throughout the cells, just happening to be easier to see I the vacuoles.  Quite what role they play remains speculation: barium is not required for plant nutrition and is, indeed, toxic in high concentrations.   It is also scarce in the soft waters where Closteriumis most often encountered, both in absolute terms and relative to other trace metals, which only adds further to the mystery.

That’s enough about Moss Dub for now.  A few words about Ennerdale Bridge, where we were based before I sign off from this post.  I usually stay at the Shepherd’s Arms when I am in the area and Keith and his staff hosted most of us and fed all of us.  It is a comfortable, unprententious inn, living mostly off walkers doing the Coast-to-Coast walk and with a menu that managed to put a smile on the faces of vegetarians and non-vegetarians alike.  Our daytime events took place in the community room of The Gather, a community-owned and run café and shop,  That gave us the satisfaction of knowing that the money we paid for the room was going to good use.   Their coffee keeps me going during long days of fieldwork in the area so I’m keen to make sure that they thrive!


Closterium lunula (400 – 663 micrometres long) from Moss Dub, showing the terminal vacuole (ringed) and (below) a close up showing rectangular crystals of barium sulphate inside the vacuole.


Brook, A. J., Fotheringham, A., Bradly, J., & Jenkins, A. (1980). Barium accumulation by desmids of the genus Closterium (Zygnemaphyceae). British Phycological Journal 15: 261-264. https://doi.org/10.1080/00071618000650251


Quekett Microscopy Club and British Phycological Society members getting stuck into analysis of samples from Moss Dub and the Ennerdale valley at The Gather, Ennerdale Bridge, September 2019.

Algae behaving selfishly …

My most recent trip to Ennerdale Water was on a wonderful windless winter day, offering perfect reflections of the snow-dusted peaks beyond the lake. It was a cold day but I was well wrapped-up and could enjoy both the long-distance views and the close-ups of nature around the lake’s margins.   One of the small streams that I crossed as I skirted the perimeter of the lake had patches of green algae growing on its submerged stones and even a quick examination showed it to be coarser than the green algae that covered most of the larger stones on the lake bed itself, as well on those in the River Ehen, just below the outfall.   When I managed to get specimens under my microscope I saw that the algae on the lake bed was Spirogyra (which I have seen here before; see “A lake of two halves”) whilst that in the inflow stream was Oedogonium.

I’ve written about Oedogonium before, and lamented the problems we face when we try to identify the species within this large genus (see “The perplexing case of the celibate alga”).   Ironically, a couple of weeks after I wrote this, I encountered a population of Oedogonium in another Cumbrian stream that did have sexual organs (see “Love and sex in a tufa-forming stream”).  However, this was the exception that proves the rule, as I have not seen a sexually-mature population of Oedogonium since.  The population I found beside Ennerdale was not sexually mature either but it did show a different, but equally effective, means of going forth and multiplying.

In the left hand diagram below we see a vegetative cell from an Oedogonium filament that has split open, allowing a vesicle to be extruded within which a single zoospore has formed.   This has a ring of flagella at one end, resembling a monk’s tonsure (you can just see these flagella in the photograph).   The other two photographs show the monk’s bald pate, though the fringe of flagella is not very clear.    The transparent vesicle swells and eventually ruptures, releasing the zoospore, which swim around for an hour or so, before settling on a new substratum and growing into new filaments.

Zoospores of Oedogonium from a stream flowing into Ennerdale Water, January 2018.   Scale bar: 25 micrometres (= 1/40th of a millimetre). 

In my material, the new filaments were mostly attached to mature Oedogonium filaments; however, this is probably partly an artefact and, in the field, they would almost certainly also settle on rocks and other surfaces too.   You can see, in the diagram below, how the “bald” end of the zoospore has started to differentiate into a holdfast that will secure the cell to the substrate whilst, over time, the other end will start to divide to produce the first cells of the new filament.  The whole process is described in a series of papers by Jeremy Pickett-Heaps (see reference list below).

Why did I see zoospore formation in this particular sample?   I don’t know for sure but it may be because I let a longer than usual time elapse between collecting and examining the sample.   This one had sat around in a cool box and fridge for four days, whereas I usually manage to check them within 24 hours.   Neglect can be a useful tool in the phycologist’s arsenal, as many freshwater algae see no need to indulge in anything more taxing than routine cell division for as long as the habitat keeps them replenished with whatever light, nutrients and other resources that they need.   Only when this is no longer the case do the algae start to channel resources into survival strategies.

Oedogonium zoospores germinating into new filaments, both epiphytic on mature filaments.   From a stream flowing into Ennerdale Water, January 2018. .   Scale bar: 25 micrometres (= 1/40th of a millimetre). 

Although I used the phrase “go forth and multiply” in an earlier paragraph, these Oedogonium cells are actually “going forth” rather than “multiplying” as the process we are watching only produces a single new cell.  However, were this zoospore to be released in a stream rather than a sample bottle, then there is a good chance that it would have been washed downstream and that a few of the many zoospores might have settled on a suitable habitat away from the constraints of their former home.   Asexual reproduction is a dispersal mechanism that results in the spread of genetically-identical copies of the parent cell.  For a sessile organism, this strategy allows a single genotype to move on from less-favourable locations and to exploit the potential of nearby locations.

The word “reproduction” is misleading as the mixing of genetic material that we associate with sex doesn’t take place.  The end product is a clone of a successful Oedogonium filament growing somewhere else.   However, taking the “sex” out of “asexual” removes a huge potential for innuendo, and readers who have battled this far through a post on nondescript green filaments deserve a reward.  So let’s finish with Woody Allen’s definition of masturbation as “sex with someone you love” and suggesting that the cytological huffing and puffing involved in zoospore production may not have the romance of sex but nor does it lead to any of the complications which result from sex either.   The alga gets offspring that are 100% identical to itself, just slightly further downstream and there is no risk of mixing with inferior genotypes.   That’s about as “selfish” as the “selfish gene” can get.


Pickett-Heaps, J. (1971).   Reproduction by zoospores in Oedogonium. I. Zoosporogenesis.   Protoplasma 72: 275-314.

Pickett-Heaps, J. (1971).   Reproduction by zoospores in Oedogonium. II. Emergence of the zoospore and the motile phase. Protoplasma 74: 149-167.

Pickett-Heaps, J. (1972).   Reproduction by zoospores in Oedogonium. III. Differentiation of the germling.  Protoplasma 74: 169-173.

Pickett-Heaps, J. (1972).   Reproduction by zoospores in Oedogonium. IV. Cell division in the germling and the possible evolution of the wall rings.   Protoplasma 74: 195-212.

See also “The River Ehen in March” for some further perspectives on asexual reproduction in algae.

View from near our sampling site on Croasdale Beck, looking towards Ennerdale Bridge, January 2018.


Ecology in the Hard Rock Café …

Before I was diverted by the delights of Bukhara and Samarkand, I was writing about the struggles that aquatic plants have to undergo in order to obtain the carbon that they need for photosynthesis (see “Concentrating on carbon …”).   In this post, I want to show the scale of the effect of inorganic carbon supply on the diatoms that we find in freshwaters.

My earlier post pointed out that aquatic plants have two possible sources of carbon to use for photosynthesis: dissolved carbon dioxide or bicarbonate.   The latter is derived, ultimately, from the rocks through which the water seeps before ending up in a stream or river.   Calcium carbonate, in turn, reacts with hydrogen ions in the water to form the bicarbonate that plants can use for photosynthesis.   A rock such as limestone, which is made of calcium carbonate, for example, provides a better supply than a hard siliceous rock such as granite.

Aquatic biologists use the term “alkalinity” to refer to the relative amounts of carbon dioxide, bicarbonate and carbonate in water.   This can confuse people as, in this context, “alkalinity” has little to do with the pH of the water itself and, indeed, water that is alkaline (i.e. has pH > 7) does not have to have a high alkalinity.   For now, just accept that low alkalinity water has little bicarbonate relative to dissolved carbon dioxide, whilst high alkalinity water has mostly bicarbonate and relatively little dissolved carbon dioxide.   In practice, alkalinity is a good indicator of the geology underlying the catchment from which a sample was collected, with low values associated regions of hard rocks (such as the Ordovician granites in Ennerdale’s catchment) and high values particularly associated with limestone and chalk.

I’ve spent a quarter of a century trying to understand how diatoms react to pollution and one of the surprising by-products of those studies is the realisation that alkalinity is just as important as pollution in determining the diatoms that are found at a site.   This is the case for most groups of freshwater organisms, but the scale of the effect on diatoms is particularly strong, as the graph below indicates.

Relationship between alkalinity and the average TDI at 430 UK river sites (r2 = 0.52).   The blue line shows a regression line fitted to the 10th percentile using the “quantreg” package in R. 

This graph shows a data from 430 UK sites where at least one TDI (Trophic Diatom Index) measurement was available, with alkalinity plotted on a logarithmic scale on the x axis and the TDI on the y axis.   There is a clear relationship between the two variables with about half of all the variation in the TDI accounted for by alkalinity (i.e. geology) alone, and this is manifest, in particular, by alkalinity setting a “floor” below which the TDI is unlikely to fall at any given alkalinity value (indicated by the blue line).  The red line, then, indicates the variation in TDI due to other factors, mostly human pressures such as eutrophication.

The blue line, in other words, indicates the best that the TDI is likely to be at any given alkalinity and if we were to look at samples which plot close to this line we will see quite marked differences in the diatoms as we moved from the low end towards the high.   When alkalinity is low, we will find Tabellaria flocculosa, some Brachysira species (e.g. B. neoexilis) and maybe a few Eunotia species too.  As alkalinity increases, so the diatom assemblage will be dominated by Achnanthidium minutissimum and relatives, but we will also see Hannaea arcus and Fragilaria gracilis, amongst other species.   We will see some Achnanthidium and Fragilaria species at low alkalinity, too, but either different to those at moderate to high alkalinity or in lower numbers.

There are several possible explanations for this but Brian Moss, in a classic paper from 1972, suggested that the availability of dissolved carbon dioxide was a major factor.  The “soft water” species, in other words, were better adapted to life without bicarbonate but were out-competed in moderate and hard water where the supply of bicarbonate was greater.   Very roughly, this switch from domination by free carbon dioxide users to bicarbonate users occurs at no more than 20 mg L-1 CaCO3.   There is more going on than just the supply of inorganic carbon: low alkalinity water is more likely to have low pH, which brings a separate set of challenges to aquatic organisms, and very high alkalinity water is often associated with productive agricultural areas.  This means that effects at both ends of the scale may be hard to separate completely from human pressures.  However, the broad story that emerges is that hard rock, in ecology as in music, is not to everyone’s taste.


Moss B. (1973).  The influence of environmental factors on the distribution of freshwater algae: an experimental study. II. The role of pH and the carbon-dioxide-bicarbonate system.  Journal of Ecology 61: 157-177.

A bigger splash …


This post continues my occasional series on insignificant black or dark brown spots on submerged stones (see “Both sides now …” for another recent episode).  I found these particular specimens on a cobble in Croasdale Beck in Cumbria, close to my regular haunts around the River Ehen and Ennerdale Water and thought that, with algae grabbing headlines for the wrong reasons yet again, I should write something positive about them.   What kind of weird world do we live in when people think it strange that algae thrive in a warm, well-lit swimming pool, whilst simultaneously lauding other people who devote four years of their lives to practising jumping into that same pool?


Colonies of Chamaesiphon cf fuscus (mostly 2-3 mm in diameter) growing on a submerged cobble in Croasdale Beck, Cumbria, August 2016. 

There was something about the regularity of the outlines of the dark brown / black spots on some of the more stable stones in this flashy beck that attracted my attention.   I’ve scraped a lot of dark smears and smudges off rocks in the past and often been disappointed when all I find are inorganic iron or manganese deposits.  Over time, one gets an eye for what is and is not an algal growth (or, for that matter, a submerged lichen) and even, in some cases, for the type of alga that formed the growth.   In this case, I had a good idea, straight away, that I was looking at a member of the genus Chamaesiphon, a cyanobacterium (blue-green alga).

Members of this genus are unicellular and form dense mats of cells that can be difficult to photograph.   I could not get a really clear view of individuals within this particular colony so, instead, have included some of Chris Carter’s photographs of another member of the genus.   You can see the short, club-shaped cells, each in a sheath and many topped by small “exospores” which bud off from the mother cell to propagate the colony.   The sheath has a brown tinge, presumably to the “sun-screen” compounds that we have met before in cyanobacteria.   Most of the members of the genus live on submerged rocks, but a few live on other algae (see “More from the River Ehen”).   Most of the rock-dwelling species indicate at least good conditions in rivers, but one species, C. polymorphus, is tolerant of more enriched conditions, which complicates use of a straight genus-level identification for rapid assessments.


Chamaesiphon polonicus from Caldbeck, Cumbria, photographed by Chris Carter.  Top left: looking down on colony; other images: side views showing cells in their sheaths and, in a few instances, with exospores. 

Oddly (to me at least) press coverage of the Olympic diving pool story has only used the word “algae”, never telling us what sort of alga is responsible for the problem.   This is equivalent to the commentators saying that “animals” have just made a perfect leap off the 3 metre springboard, leaving the audience to work out whether the subsequent splash was made by a slug or a human.

But I should end on a positive note: better, perhaps to compare the algae not with the divers but with the judges who assign the final scores.   That’s because a few minutes mooching around a stream or beside a lake can usually reveal enough from the types of algae that live there to give some insights into the health of the stream.   My visits to Croasdale Beck over the past year or so have shown me enough to suggest that this little Cumbrian stream probably deserves the algal equivalent of an Olympic medal.  But I doubt that we’ll get much press coverage for saying that…