Shuffling the pack


The next group of algae I’m going to consider in my review of higher taxonomy and systematics, the Cyanobacteria, or “blue-green algae”, present some significant challenges.  Not least of these is a shift over the past forty years from being classified according to the rules of botanical nomenclature to being classified according to the International Code of Nomenclature of Bacteria.  The former assumed species could be defined from field material on the basis of morphology, with “type specimens” preserved in herbaria; the latter uses axenic (i.e. pure) cultures as the basic taxonomic unit, and allows a wider range of attributes than just morphology.   In recent years, as those who follow this blog will know, properties such as gene sequences have also been used to define species although the Code of Botanical (now Biological) Nomenclature still requires a description of the any new species that are described, with an expectation that the morphology will take a prominent role in that description.  As this post will show, morphology is no longer such a reliable indication of how Cyanobacteria are organised as it was in the past.

For practical purposes, many Cyanobacteria fall into the same size range as other algae, live in communities that include many protist groups and can be identified using similar techniques as would be employed to identify other algae.  They also have a form of photosynthesis that produces oxygen as an exhaust gas, in contrast to other bacteria which are capable of photosynthesis. This means that a default view of the Cyanobacteria as “algae” is a reasonable starting point for a field ecologist.   However, at an intercellular scale, the Cyanobacteria are very different to other algae, and we should never lose sight of the fact that they actually belong to a different Domain to other algae.

The problems are clear when I compare the morphology-based classification that I used when I first taught classes on algae in 1990 with the classifications that are accepted now.  Then, Cyanobacteria were divided into three or four orders, typically:

  • Chroococcales – single cells or cells loosely-bound into irregular gelatinous colonies
  • Oscillatoriales – filamentous forms lacking heterocysts
  • Nostocales – filamentous forms with heterocysts

The high-level classification, in other words, was based solely on whether or not the organism formed filaments and, if so, whether or not it possessed heterocysts (specialised cells responsible for nitrogen fixation).  This made logical sense when your primary source of insight is morphology.  Unfortunately, more recent studies have shown that it bears little relationship to the genetic relationships amongst the organisms that have been revealed over the past thirty years or so.   A more recent organisation is given in the diagram below.

First, note that this shows subclasses, rather than orders, within the class “Cyanophyceae” (the only class in the division Cyanophyta).   There is rarely unanimity amongst experts on the appropriate organisation of high-level classifications so just bear with me on this one.   Of the four sub-classes, one, Nostocophycidae, contains a single order (Nostocales) which includes all the heterocyst-bearing forms.  No change there.   However, the other two classes diverge very much from the older classifications in that they both contain a mixture of filamentous and non-filamentous forms.


The organisation of the Cyanobacteria (blue-green algae) division into four sub-classes.  Filled boxes indicates the classes that are represented in UK and Irish freshwaters.   Organisation follows Algaebase.   The image at the top of this post shows a Microcystis bloom at Ladybower Reservoir (photo: Chris Carter)

The Oscillatoriophycidae is a good example, with five sub-classes, four of which are represented in the UK and Ireland.  Two of these have featured in several posts (see Appendix) so you can see for yourself just how different they are in appearance.  The Oscillatoriales includes filamentous forms without heterocysts whilst the Chrococcales has taxa that either exist as single cells or in masses loosely-bound within gelatinous colonies.    A similar situation exists within the Synechococcophycidae; indeed, some genera that would formerly have been considered to be relatives of taxa within Oscillatoriales (e.g. Schizothrix and Heteroleibiana) are now included in families in this group.   There is, however, still more work to be done to unravel all the relationships within this sub-class.   The current understanding is that there is a single order (“Synechococcales”) but a great number of families.  Similarly, all the heterocystous forms are grouped into a single order, the Nostocales, within the Nostocophycidae, also divided into a large number of families.


Organisation of the Oscillatoriophycidae showing the orders that include genera found in UK and Irish freshwaters.  

I always stress that taxonomy and identification are two distinct crafts: the taxonomist calls on a wide range of tools to find natural groupings of species at different levels whilst an ecologist only needs a parsimonious route to an unambiguous identification.  For the purposes of identification, recognising whether an organism is filamentous or not is a logical early step, even though both options will contain representatives of both Oscillatoriophycidae and Synecchococcophycidae.   We need to recognise that some of the characteristics that contribute to our taxonomic understanding (gene sequences, arrangement of thylakoids) are useless from the point of view of someone trying to name an organism encountered in a field sample but, at the same time, the taxonomist’s standpoint will not necessarily capture all of the features that explain how an organism contributes to energy and nutrient flow within ecosystems.


Calothrix stagnalis: a member of the Nostocales.  Note the heterocysts at the base of the filaments (photo: Chris Carter)


Mai, T., Johansen, J.R., Pietrasiak, N., Bohunciká, M. & Martin, M.P. (2018).  Revision of the Synechococcales (Cyanobacteria) through recognition of four families including Oculatellaceae fam. nov. and Trichocoleraceae fam. nov. and six new genera containing 14 species.  Phytotaxa 365: 1-59.

Palinska, K.A. & Surosz, W. (2014).  Taxonomy of cyanobacteria: a contribution to consensus approach.  Hydrobiologia 740: 1-11.


Links to posts describing representatives of the major groups of Cyanobacteria found in freshwaters.  Only the most recent posts are included, but these should contain links to older posts (you can also use the WordPress search engine to find older posts).

Group Link
Synechococcales Chamaesiphon: A bigger splash

Heteroleibleinia: River Ehen … again

Chrococcales Aphanothece: No excuse for not swimming …

Gloeocapsa: The mysteries of Clapham Junction …

Oscillatoriales Microcoleus: How to make an ecosystem

Oscillatoria: Transitory phenomena …

Phormidium: In which the spirit of Jeremy Clarkson is evoked …

Pleurocapsales Watch this space …
Spirulinales Spirulina/Arthospira: Twisted tales …
Nostocales Nostoc: How to make an ecosystem (2)

Rivularia: Both sides now

Scytonema, Stigonema, Tolypothrix: Tales from the splash zone

Some other highlights from this week:

Wrote this whilst listening to: Leonard Cohen’s posthumous album Thanks for the Dance.   And, as I used it to name a post, Joni Mitchell’s Both Sides Now.

Cultural highlights:  David Hockney: Drawing from Life at the National Portrait Gallery.   Great examination of the importance of drawing and observation to artistic practice.   By coincidence, another post I’ve cited is named after one of Hockney’s paintings

Currently reading: Robin Wall Kimmerer: Gathering Moss (Oregon State University Press).  A collection of essays on the natural and cultural history of mosses.

Culinary highlight: dinner at The Sichuan on City Road in London.


How to make an ecosystem (2)


My most recent visit to Ennerdale and the River Ehen almost did not happen: unexpected overnight snowfall led to my wheels spinning on the Whinlatter Pass before I retraced my steps to Braithwaite and followed roads at lower altitudes around the outskirts of the fells.   Fieldwork in the morning took place amidst intermittent snow showers but, by the afternoon, it was dry if not quite as balmy as the visit I described in “Croasdale Beck in February”.   “Unseasonable”, I was reminded, is a two-edged term.

There was little incentive to linger with my arm in the agonisingly cold water, so this post is about some algae growing on dry land that caught my eye.   Amidst the gravel in a farmyard in Ennerdale Bridge I saw some dark brown leathery growths that I recognised straight away as the Cyanobacterium Nostoc commune (see “Nosing around for blue-green algae …”).  It looks rather nondescript, even slightly unsavoury, with the naked eye but, under the microscope, the rosary-like structure of the filaments suspended within a jelly-like matrix is revealed.  The slightly larger cells with thicker walls and lighter contents are the heterocysts, responsible for fixing nitrogen from the atmosphere (fulfilling the same function as the nodules on the roots of legumes).


A patch of Nostoc communein a farmyard in Ennerdale Bridge in April 2019.   The picture frame covers about 30 centimetres. 


Nostoc communefrom Ennerdale Bridge under the microscope.  Scale bar: 10 micrometres (= 100thof a millimetre). 

This type of coarse, well-drained gravel is a good habitat for Nostoc and, once you know what you are looking for, it is a common sight on gravel driveways, usually to the annoyance of the owners.   If there is only a small amount, the best way to control it is simply to pick up the colonies and toss them onto the compost heap.  However, once it is established, this can be a big undertaking and many people are quite happy to tolerate some of this brown gunk on their driveways.   On the other hand, it can sometimes get out of hand and the consequences of not doing anything are well illustrated by the photograph below.  The Nostoc colonies have spread but these, in turn, have created a habitat into which first mosses and later grasses can establish.

This small farmyard on the edge of the Lake District contains, in short, the first stage of an ecological succession.  We could think of a gravel driveway as a mini-desert, as the copious Cumbrian rainfall will not be retained in the surface layers, making it hard for plants to survive.   However, if a tough organism such as Nostoc is able to establish itself, then this, in turn, will trap water and make the driveway more amenable to slightly more fussy organisms such as mosses.   As the moss and Nostoc grow together so, eventually, grasses are able to establish too.  Were there to be no interruption to this process then, eventually, decades later, we might even see trees growing on this driveway.

It is hard to imagine, but just about every type of terrestrial habitat started out, aeons ago, as a bare rock surface.  Various forms of physical weathering start the process of breaking this up allowing, over time, organisms such as Nostocto get a foothold and convert the virgin surface into a mature ecosystem (you can read about another example in “How to make an ecosystem”).   It may take centuries for this to happen in the natural world, so it is particularly fortuitous to see this human-assisted succession so well developed.  At some stage, I suspect, the owner will decide that enough is enough, and rake the gravel.  Meanwhile, however, we have a rare opportunity to reflect on the role that primitive micro-organisms play in shaping even the grandest of our natural habitats.


A lawn of Nostoc, moss and grass growing on a gravel driveway in Ennerdale Bridge, April 2019.  


Miles, J. & Walton, D.W.H. (1993).  Primary Succession on Land.  Special Publication of the British Ecological Society 12, Blackwell Scientific Publications, Oxford.

More about measuring biomass …

The previous post showed how the proportions of green algae and diatoms changed as the total quantity of algae in the River Ehen waxed and waned over the course of a year.   The BenthoTorch, however, also measures “blue-green algae” and so let’s look at how this group changes in order to complete the picture.

Before starting, though, we need to consider one of the major flaws of the BenthoTorch: its algorithms purport to evaluate the quantities of three major groups of algae yet, in my posts about the River Ehen I have also talked about a fourth group, the red algae, or Rhodophyta (most recently in “The only way is up …”).  Having pointed a BenthoTorch at numerous stones with thick growths of Audouinella,we can report that Rhodophyta seem to be bundled in with the blue-green alga signal, which is no great surprise given the similarity in their pigments.  It is, however, one of a number of examples of the need to interpret any BenthoTorch results with your brain fully engaged, and not just to treat outputs at face value. Similar questions need to be asked of the Xanthophyta and Chrysophyta, though the latter tend not to be common in UK streams.


Relationship between the proportion of “blue-green algae” (Cyanobacteria and Rhodophyta) and the total quantity of benthic algae (expressed as chlorophyll concentration) in the River Ehen (c.) and Croasdale Beck (d.).  The blue lines show quantile regression fits at p = 0.8, 0.5 and 0.2.  

In contrast to the green algae and diatoms, the Cyanobacteria/Rhodophyta signal shows a strong negative relationship as biomass increases though, again, there is enough scatter in this relationship to make it necessary to approach this graph with caution.  I suspect, for example, that the data points on the upper right side of the data cloud represents samples rich in Audouinella, which tends to occur in winter when biomass, generally, is much greater.   On the other hand, Croasdale Beck, in particular, has a lot of encrusting Chamaesiphon fuscus colonies which are pretty much perennial (see “a bigger splash …”) but whose relative importance in the BenthoTorch output will be greatest when the other two groups of algae are sparse.   I suspect that encrusting members of this genus are favoured by conditions that do not allow a high biomass of other algae to develop, as these will reduce the amount of light that the Chamaesiphonreceives.

Thicker biofilms in the River Ehen often have some narrow Phormidium-type filaments as well as small bundles of nitrogen-fixing Calothrix, but the overall proportion is generally low relative to the mass of diatoms and green algae that predominate.    But that is not really telling us the whole story.  I finished my previous post with a graph showing how the variation in biomass increases as the biomass increases.  The heterogeneity of stream algal communities, however, cannot be captured fully at the spatial scale at which the BenthoTorch works: there is a patchiness that is apparent to the naked eye: one of our sites has distinct mats of Phormidium autumnale towards one margin, and dense Lemaneagrowths in the fastest-flowing sections, largely attached to unmovable boulders, which makes biomass measurement very difficult. I’ve also written about distinct growths of Tolypothrix and its epiphytes (see “River Ehen … again”), another alga which forms discrete colonies at a few locations. I try to collect a random sample of stones from a site but there are constraints, including accessibility, especially when the river rises above base flow.   In the River Ehen we also have to take care not to disturb any mussels whilst removing stones.

Whilst our sampling cannot really be described as “random” I do think that there is sufficient consistency in the patterns we see for the results to be meaningful. We could spend a lot more time finessing the sampling design yet for little extra scientific gain.   I prefer to think of these measurements as one part of a complex jigsaw that is slowly revealing the interactions between the constituents of the dynamic ecosystem of the River Ehen.   The important thing is to not place too much faith in any single strand of evidence, and to have enough awareness of the broader biology of the stream to read beyond the face value indications.

Both sides now …


I diverted from my usual haunts in the upper River Ehen in Cumbria recently in order to explore Ennerdale Water in greater detail.   I am used to see it from the western end as we do our fieldwork, but the length of the journey to and from the River Ehen means that we rarely have time to linger.  Finally, however, we found a July day when we could circumnavigate the lake.  “July day”,”Lake District” and “fieldwork” sounds like an intoxicating combination.  However, the photograph above shows it was not quite as idyllic as it might have been (or, even, as it was on the day before).  Hence the title of this post, borrowed from a beautiful Joni Mitchell song which includes the line “But clouds got in my way”.

In the far past, the lakes of the Lake District were thought to have “evolved” at different speeds following their formation at the end of the last Ice Age.  Ennerdale Water and Wastwater, surrounded by hard volcanic rocks which erode very slowly, were regarded as the two most “primitive” lakes, whilst Windermere and Esthwaite Water were thought to be the two most “evolved”.   That is now known not to be the case: the geology is very important in determining the type of lake, not just because erosion is the source of the inorganic salts that give the water a particular chemical character, but also because this influences how man uses the lake.   In the case of Ennerdale Water, only about five per cent of the catchment is cultivatable, and this, in turn, influences the amount of inorganic fertiliser that is added to the meagre supply of salts provided by the underlying rocks.   Ennerdale is, as a result, one of the least chemically-disturbed of all English lakes.

At the far south east end of the lake, close to where the River Liza enters the lake, I was intrigued to see some very dark spots on the rocks.  They looked like they might be cyanobacterial colonies, so I picked a few off with my forceps and put them into a bottle for later investigation.  When I was able to look at them, the following day, I saw clumps with brown filaments radiating out, and each gradually narrowing towards the tip.   Closer examination showed that the cells that made up each filament had a blue-green colour, but were each enclosed in a brown pigmented sheath.   The filaments showed a characteristic form of “false” branching, in which the daughter filament breaks off from the mother, but is contained within the same sheath.   At the base of many of the filaments, I could see a modified cell (slightly lighter and less granular than the others) called a “heterocyst”, which was responsible for nitrogen-fixation.   These are all characteristics of the genus Rivularia, which is a good indicator of very high status water.


A cobble (about 15 cm long) from the littoral zone of Ennerdale Water, SW end.  The dark cyanobacterial colonies are about 3-4 mm across.


Rivularia biasolettiana from the littoral zone at the south east end of Ennerdale Water, Cumbria, UK.  a. low power (x100) image showing radiating filaments gradually narrowing in width; b. filaments showing false branching (a heterocyst is present, but hard to see); c. false branching in a filament of Rivularia with heterocyst arrowed.   Scale bar (b. and c.): 10 micrometres (= 1/100th of a millimetre).

The presence of Rivularia here was interesting to me for several reasons.   First, it continues a series of observations that suggest that this genus is not confined to hard water habitats in Britain and Ireland, as once thought (see “more about Rivularia” and links) although earlier posts have also referred to its presence in soft water habitats in Norway.   I’m also fairly sure that the organism that I collected from this trip to Ennerdale is different to the one that I find in the River Ehen and, indeed, at other locations around the Ennerdale perimeter, but that is a subject for another day.

The second comment to make is that the presence of cyanobacteria (blue-green algae) is usually a sign of an unhealthy, enriched habitat, not the very high quality habitat that Ennerdale, in fact, represents.  Blue-green algae in the phytoplankton is, very often, a bad sign, suggesting enrichment by inorganic nutrients.  Several species combine the ability to fix nitrogen with positive buoyancy, which means that they have two distinct advantages over other algae as they compete to exploit limited light and nutrients.  Some of these blue-green algae are also toxic, which has implications for how the lake and its water is used.

Those blue-green algae that live attached to surfaces in streams and in the littoral zone of lakes play by a different set of rules, however.  As they cannot use positive buoyancy to compete for light, they are more likely to be overgrown by faster-growing algae in the scrabble to capture available light.   This means that investment in expensive nitrogen fixation machinery is only an advantage when other algae, too, are very short of nutrients.  And a shortage of nutrients is the natural state for most freshwater ecosystems.

I chose the title of this piece, originally, because I felt that I had looked at Ennerdale Water from both sides now.  However, the same lyric could just as well apply to blue-green algae.   The water manager sees them as a problem; in some situations, however, they can be a positive sign.   So we can, in fact, look at blue-green algae from both sides too … from give and take and still somehow … I really don’t know life (microscopic life especially) at all ….

Poking around amongst sheep’s droppings …

A couple of kilometres away from the stream featured in the previous post is an old quarry that we visit each year as part of this course (see “Nosing around for blue-green algae …”).   In a damp flush at the edge of the quarry floor, we found some patches of what looked, from a distance, like sheep droppings.   A useful strategy, shared by most of the human race, is to assume that anything that looks like a sheep’s dropping probably is a sheep’s dropping, and not to go prodding at this with a finger.   However, a curious soul in the distant past with a sense of adventure ignored this precept and discovered that a few of these were, in fact, growths of Cyanobacteria.   Most natural historians, wisely, focus their attention on more spectacular aspects of life on earth; however, a few of us have retained this childish instinct to poke at anything that looks like sheep’s droppings.


Scytonema sp. from a flush at Whitbarrow Quarry, Cumbria, May 2015

A small part of one of these growths, teased out and mounted on a cover slip, reveals itself to have the characteristics of the genus Scytonema although, today and despite a long hunt, I could not find any filaments that presented themselves in a suitable position to photograph. The illustration below, therefore, is of a growth of Scytonema from another calcareous site in Cumbria. The Cyanobacteria is, you may remember, the modern name for the “blue-green algae” which is often confusing as many Cyanobacteria are not blue-green in colour.   What we can see here is a chain of cells (a “trichome”) which are surrounded by a thick sheath (“trichome” plus “sheath” equals “filament”, in Cyanobacteriological lore).   The sheath is a yellow-brown colour, due to a pigment called “scytonemin” which acts as a sunscreen, absorbing ultra-violet radiation and, in the process, obscuring the blue-green colour of the trichome within.


Scytonema sp. from a calcareous flush at Sunbiggin Tarn, May 2005.   Scale bar: 10 micrometres (= 1/100th of a millimetre).

Two other characteristics of Scytonema are visible in the photograph.   Both the left and right hand pictures shows “false branches”: if the trichome breaks for any reason, either or both of the broken ends keep growing and break out of the filament. The left hand illustration is a single false branch and, just to the left of the branch you can see the distinct “heterocyst”, a cell where nitrogen fixation takes place.   The wall of the heterocyst is thicker than that of other cells, as nitrogen fixation can only occur in the absence of oxygen.

Walking back from the quarry towards the minibus, Allan pointed up at dark patches on the cliffs looming over us.   It was Gloeocapsa alpina, the same species that we met a short while ago in a cave on Malta (see “The mysteries of Clapham Junction …”).   The cliffs at Whitbarrow are, in effect, a vertical “desert” from the point of view of any organism that aspires to live there. These Cyanobacteria, with their ability to “re-boot” on those occasions when conditions are favourable for growth, have an advantage here.   One guesses that the damp climate of north-west England is slightly more forgiving than that of a Mediterranean hillside but it is still a tough habitat in which to survive.


Growths of Gloeocapsa alpina (arrowed) on the cliffs at Whitbarrow Quarry, May 2015.

The origins of life …

I have been reading Richard Fortey’s book “Survivors” (see “”They don’t do much, do they?””) which is a rare and precious thing in the world of natural history writing as he devotes two whole chapters to the algae.   The first of these describes his encounters with living stromatolites in Shark Bay, in Western Australia.   The point he is making as he writes is that he did not just make an extraordinarily long journey to get there (first, get to Perth, then travel a further 800 km north …) but that he is also making a similarly long journey back in time, as stromatolites are survivors from the Precambrian era and, indeed, may have played a vital role in creating the oxygen-rich atmosphere that we take for granted.

I have a polished specimen of a stromatolite, purchased from a fossil shop in Durham, which shows the characteristic fine laminations. Each of these represents a layer of Cyanobacteria (blue-green algae) filaments which have, in turn, trapped sediment particles.   The laminations are, in turn, formed into dome-like structures, reflecting the vertical growth of the filaments in search of sunlight.   This stromatolite is from the Ordovician era (I think), which dates it to between 443 and 485 million years ago.


A stromatolite from Argentina, possibly from the Ordovician era. The specimen is 10 cm long.

Not only were stromatolites extremely abundant in the Precambrian (approx. 4.6 billion to 540 million years ago) but the organisms from which they were formed appear to be very similar to Cyanobacteria that can still be found today.   They are extremely delicate structures that thrived, at least in part, because there were few other organisms in the Precambrian that could compete with them or graze them and, consequently, had shallow marine habitats to themselves for an extremely long time. During this period, they were busily photosynthesising away, taking carbon dioxide and water and converting it to simple sugars (which they needed to grow) and oxygen (which was, as far as the Cyanobacteria was concerned, a waste product). This oxygen was released as tiny bubbles (see “Ecological yin and yang…”) which, ever so slowly, accumulated in the atmosphere.   Maybe it is no surprise that the Precambrian, the era before fossils of multicellular organisms are common, lasts for about four fifths of the entire lifespan of the earth: it took this long for all those tiny bubbles to add up to enough oxygen to allow more complicated organisms to survive.

And what did those multicellular organisms feed upon?   That’s right: the Cyanobacteria had sown the seeds of their own destruction.   There is evidence not just of a gradual decline of stromatolites through the later Precambrian and into the Cambrian and Ordovician eras, but also of a resurgence of stromatolites after the mass extinction at the end of the Ordovician (which would have removed the multicellular grazers and left our tough little Cyanobacteria behind).   Stromatolites are found sporadically throughout the fossil record and in a small number of locations in the present day, but their heyday lies far in the past.


Bengston, S. (2002). The early worm catches the – what?   Pp. 289-317. In: The Fossil Record of Predation (edited by Kowalewski, M., and Kelley, P.H.). The Paleontological Society Papers 8, The Paleontological Society, Boulder, Colorado.

Fortey, R. (2011).   Survivors: the Animals and Plants that Time Has Left Behind.   Harper Press, London.

Sheehan, P.M. & Harris, M.T. (2004). Microbialite resurgence after the Late Ordovician extinction. Nature (London) 430: 75-78.

“They don’t do much, do they?”

I spent the early part of yesterday evening listening to Richard Fortey talk on “Survivors: the animals and plants that time has left behind” at Van Mildert College in Durham.   Fortey is a palaeontologist, best known for his work on trilobites, but who has also presented popular television series on natural history topics.   In this lecture, he talked about the small number of organisms that were well represented in the fossil record (some pre-dating dinosaurs) yet which had survived the various mass extinctions and which could still be found today. Did these organisms have anything in common, he wondered.

One group of organisms that he talked about were the blue-green algae (Cyanobacteria) responsible for forming “stromatolites” which are found both in Precambrian rocks over a billion years old and in a few locations today.   I’ve talked about blue-green algae in many of my posts (e.g. “More reflections from the dawn of time …”) but what amused me this evening was Richard Fortey’s anecdote about a discussion with a BBC producer as they devised a television series based on his book:

Fortey: “These are the most important organisms in the history of the earth” (commenting on their role in creating the oxygen-rich atmosphere that every organism since has relied upon)

BBC producer: “they don’t do much, do they?”

It is a subject that I have addressed several times in this blog (see “The sum of things …”, “Every second breath …”): how do we address the imbalance in natural history broadcasting between the charismatic “few” and the unfashionable “many”, bearing in mind, of course, that these are, in many cases, the hard-working, unglamorous “back office staff” who keep our planet running.   Television natural history programmers are in the entertainment business first and I’m not going to pretend that a stromatolite made of blue-green algae is, on its own, a recipe for compelling television.   But I also feel that there are possibilities that could be explored, and that we may be held back by a lack of imagination on the part of broadcasting creatives and commissioning editors.   They, too, are children of the television age, brought up on a style of broadcasting from Zoo Quest through Life On Earth and onwards that is interested only in the televisual aspects of natural history. Whoa .. hold it there … I’m coming dangerously close to criticising David Attenborough … Saint David … that’s close to heresy.


Fortey, R. (2011). Survivors: the animals and plants that time has left behind.   Harper Collins, London.

Rewriting history at Talkin Tarn …

One of the joys of modern technology is the ability to right obvious wrongs.   So, having watched my wife’s crew struggle in a regatta at Talkin Tarn, near Brampton, last weekend, having been entered in the wrong class and the wrong age category and then racing in a men’s rather than a woman’s boat, I had no option but to turn to Photoshop for justice.   Here, in a rewritten version of the Talkin Tarn regatta, you can see the DARC novice women’s coxed four half a length head of their opposition with about 400 metres to go.   The backdrop is the Pennine fells, looking north towards Hadrian’s Wall.


Talkin Tarn regatta 2014: DARC novice women’s coxed four half a length ahead of the opposition.  

I have to confess that the entertainment value of watching one crew busting a gut to go slightly faster than another crew pales after the first hour, so I wandered off to poke around the lake’s margins.   The water here contained some just discernable green specks, all between about half a millimetre and a millimetre across.   Under the microscope, these resolve into spherical colonies each composed of filaments which gradually taper from the base.   These colonies belong to a cyanobacterium (blue-green alga) called Gloeotrichia echinulata. I’ve written about relatives of Gloeotrichia in previous posts (“Blue skies and blue flowers in Upper Teesdale”, “”Looking” is not the same as “seeing” …”, “More about Rivularia”) and have commented that the large light coloured cells, called “heterocysts” are responsible for nitrogen fixation, which enables the alga to grow even when nitrogen is relatively scarce. In the case of Talkin Tarn, the scarcity is probably not absolute but relative to other nutrients such as phosphorus. It means that the Gloeotrichia can grow even when the supply of this important nutrient has run out.


Colonies of Gloeotrichia echinulata in a drop of water from Talkin Tarn, July 2014.   The drop is 12 millimetres across.

One other difference between the filaments of Gloeotrichia and those of Rivularia is the dark specks inside the cells.   These are gas vacuoles which make the cells and colonies of Gloeotrichia more buoyant than other algae. Leave a sample from Talkin Tarn to stand for a while and the colonies of Gloeotrichia will rise to the surface. This gives cyanobacteria a competitive advantage over other algae in lakes as they now have an adaptation that keeps them closer to the sunlight which powers their photosynthesis as well as one that helps them to overcome nitrogen limitation and an adaptation.   It is no surprise that cyanobacteria are so prolific in lakes around the world.


Microscopic views of a Gloeotrichia echinulata colony from Talkin Tarn.   The left hand image shows an entire colony, about half a millimetre across.   The right hand image shows the same colony at higher magnification.   Note the large, round light-coloured heterocysts at the base of each filament. Scale bar: 10 micrometres (= 1/100th of a millimetre).

Gloeotrichia also has the ability to produce compounds called “microcystins” which are powerful toxins.   There are many records of livestock, dogs and humans suffering liver damage through ingesting contaminated water.   Although the rowers at Talkin Tarn were warned to wash their hands after racing, the risk is only high if a large amount of Gloeotrichia-containing water is swallowed. That is unlikely in a sport such as rowing, where the idea is to stay in the boat, but swimming in Talkin Tarn would not be a good idea. Rowing, though, is not without such dangers: the Durham women’s novice crew’s previous opponents suffered an “ejector crab” at the start which dumped one of them into the River Wear. On that occasion, cyanobacteria in the River Wear were the least of their worries and the Durham crew were able to cruise serenely down the river to a glorious victory.   Photoshop not required.


Nosing around for blue-green algae …

One of the ironies of teaching a course on algal identification in the Lake District is that we actually take the students out of the Lake District on the first field trip in order to introduce them to the enormous variety of Cyanobacteria (blue-green algae).   This is because the southern part of the Lake District, where the FBA is located, is situated on the Silurian Slates, which means that the streams, lakes and tarns have fairly soft water.   Cyanobacteria, on the other hand, tend to be most abundant and diverse in hard water, so we drive about 40 minutes south and east from Windermere to a limestone escarpment called Whitbarrow, where there are a number of calcareous flushes and springs that are ideal for our purposes.

We always visit the floor of an abandoned quarry in this area which has several such flushes. The quarry owners had systematically removed the limestone until they had reached the Silurian Slate underneath. This, in turn, formed an impermeable layer that intercepted any water that had percolated through the limestone. The quarry floor was, typically, slippery with calcium-rich water that had seeped out from the surrounding limestone, as well as the mucilage that the algae produced.   There were also, dotted around, several unprepossessing brown objects that, to the untrained eye, could easily be mistaken for the droppings of a small animal or bird (a Peregrine falcon was circling overhead during our visit). Once the students have the courage to pick these up, they see that they are composed of a firm jelly-like substance that is, we persuade them, actually an alga and should, therefore, be dropped into one of their specimen tubes to take back for closer investigation.Nostoc_Whitbarrow_May2014

Animal, vegetable or mineral?   Colonies of Nostoc commune on the floor of Whitbarrow Quarry, May 2014.

Once back in the FBA’s laboratory (complete with panoramic views of Windermere), we can dissect out small pieces of the jelly-like material and squash it onto a microscope slide.   What they see when they peer down their microscopes is a plethora of chains of bead-like cells of a Cyanobacterium called Nostoc commune.   Most of the cells have contents that have a granular appearance, with a background of bluish-green photosynthetic pigments.   A few of the cells, however, are rounder and clearer: these are the “heterocysts”, cells that are especially adapted to “fix” atmospheric nitrogen and so help the organism survive in nutrient-poor habitats.   The jelly-like matrix slows the rate at which water evaporates from the colonies, with the outer layers drying to form a tough, leathery skin around the colony.


Nostoc commune from Whitbarrow Quarry under the microscope.   The cells are approximately 5 micrometres (1/200th of a millimetre) across.<br

There is a fascinating short paper by Malcolm Potts on the origin of the name “Nostoc”.   Because Nostoc colonies often appeared very quickly following heavy rain (because the dried colonies absorb water quickly and swell in size), there was a belief in Medieval times that Nostoc fell from the sky.   A German mystic and alchemist, Parselus, was the first to use the name “Nostoc”, claiming that it was “…excrement blown from the nostrils of some rheumatick planet.   The name, indeed, is strongly suggestive of both the Old English word Nosthryl and the German term nasenloch both, as Potts politely explains, “…that part of the human anatomy intimately associated with extracellular polysaccharide.”


Potts, M. (1997). “Etymology of the Genus Name Nostoc (Cyanobacteria)” (pdf). International Journal of Systematic Bacteriology 47 (2): 584. doi:10.1099/00207713-47-2-584

More about Rivularia

My post on Rivularia from softwater habitats (‘“Looking’ is not the same as seeing”’) prompted an email from Bryan Kennedy in Ireland with some pictures of Rivularia from a moorland stream in Co. Mayo in the west of Ireland, once again from a catchment completely lacking limestone.   Bryan estimates the calcium concentration in the water to be between 5 and 10 milligrams per litre, which means that the water here is very soft.  He also comments that it was recorded in the 1970s from the Caragh catchment in south-west Ireland (average calcium concentration: 2.15 milligrams per litre: see Heuff & Horkan, 1984).


A tributary of the Yellow River, Co. Mayo, Ireland (left) with dark brown / black colonies of Rivularia beccariana on a submerged stone (right).  Photos: Bryan Kennedy.

The photomicrographs show the colony structure very well with filaments radiating out from the centre.   The major difference between these and the Rivularia biasolettiana I photographed in Upper Teesdale (“Blue skies and blue flowers in Upper Teesdale”) is that colonies of the latter contain calcite crystals, though these were not visible in my images.  The right hand image shows the structure of Rivularia filaments very clearly; the tapering blue-green filament gradually narrowing to a colourless hair.   Note the colourless cell at the base of the filament.  This is the “heterocyst”, and is the location where nitrogen fixation takes place.  This is a very useful adaptation in the nutrient-poor habitats where Rivularia is found, as it means that, like peas and beans, it can capture nitrogen directly from the atmosphere, rather than relying upon dissolved minerals.

Nitrogen-fixation, however, needs a lot of energy and organisms do not fix nitrogen if there is a ready supply available from other sources.   Once nitrogen is abundant, species such as Riviularia are at a competitive disadvantage and it is no surprise that Rivularia and it’s close relatives are found only in remote parts of the country, given the extent to which nitrate fertiliser washes off the land and into streams and rivers.   Even in upland areas, there are often nitrogen compounds in rain water, much of it originating in the exhaust emissions from our cars.   One wonders if Rivularia might have been much more widespread a hundred years ago than is the case now.


A close-up of a Rivularia beccarina colony from the tributary of the Yellow River, Co. Mayo, Ireland.  Photos: Bryan Kennedy.  


Heuff, H. & Horkan, K. (1984).  Caragh.  Pp. 363-384.  In: Ecology of European Rivers (edited by B.A. Whitton).   Blackwell Scientific Publications, Oxford.