Life in the colonies …

Another outcome of my visit to Ennerdale Water a couple of weeks ago in July (see “Life in the Deep Zone”) was some tiny green spheres in the sample I collected from one of the small streams flowing into the lake’s north-west corner.   The stream was very short, little more than a seepage arising from a wet rush-dominated area of a field just twenty metres or so from the lake margin and, at the point which I sampled, there was a tangle of filamentous algae (Stigeoclonium, Mirsrospora and Mougeotia) as well as a distinct diatom-dominated film on exposed stones.    The colonies looked like tiny peas in my sample tray but I suspect that they were attached to rocks or aquatic vegetation before I disrupted them. Under the microscope, these turned out to be colonies of the green alga Chaetophora pisiformis, a relative of Draparnaldia and Stigeoclonium, both of which I have written about before (see “The exception that proves the rule …” and “A day out in Weardale …”.  Like those, Chaetophora has branched filaments but they differ in forming well-defined colonies that are visible to the naked eye.

The pictures below show the form of colonies very clearly.  Chaetophora colonies are firm to the touch and cannot easily be squashed under a coverslip.   I overcame this by using a cavity slide, and taking one of the smallest colonies that I could find in order to photograph it with as little damage as possible.  Note how there is a very clear edge to the colony, whereas Draparnaldia and Stigeoclonium have a mass of filaments and mucilage but no obvious border between the “colony” and the surrounding environment.  Draparnaldia sometimes forms discrete colonies (see “The exception that proves the rule …”) but these are much softer and more easily squashed onto a slide.

Top: colonies of Chaetophora pisiformis from a small stream flowing into Ennerdale Water, with a one cent coin for scale; bottom left: lower power (x40) view of a colony.  The picture frame is about two millimetres across; bottom right: medium power (x100) view of the same colony.

Viewed at higher magnifications, the branches of the filaments are clear. They tend to be clustered towards the tops of the filaments and, in this case at least, end abruptly, rather than tapering to fine hairs.  I explained in the posts mentioned above how these fine hairs are used by the algae as means of capturing the nutrients that they need.  Chaetophora can form these hairs, but it does so less often, in my experience, than Draparnaldia and Stigeoclonium.   There will be dead and decaying vegetation in the rush-dominated swamp from which the stream originates, and the enzymes that these algae produce will be able to harvest any phosphorus from organic particles that result from this decay.  That’s the theory for Stigeoclonium at least, but I suspect that the colonies of Chaetophora are also highly efficient recycling units: the filaments are embedded in a firm mucilage that is far more than an inert polysaccharide gunk.   Any phosphorus that is released from a filament will be far more likely to be hoovered up by another filament than to drift downstream whilst the phosphatase enzymes will also be on hand at the colony surface to savenge any stray nutrients from the seepage.  These tight colonial forms are, in other words, fortresses of plenty in an otherwise inhospitable landscape: well adapted to nutrient-stressed situations and, as a paucity of nutrients is the natural condition of streams, the presence of these colonies is a good sign that this stream is in good condition.

Filaments of Chaetophora pisiformis from a small stream flowing into Ennerdale Water, July 2018.  Scale bar: 20 micrometres (= 1/50th of a millimetre). 


Whitton, B.A. (1988).  Hairs in eukaryotic algae.   pp. 446-480.  In: Algae and the Aquatic Environment (edited by F.E. Round).  Biopress, Bristol.


Life in the deep zone …

The view above – looking along Ennerdale Water from the western end – is one that I’ve used before in this blog.  The difference today is that there is about twenty metres of foreshore exposed.  Normally, water covers all the area in the foreground.   Not today: Ennerdale Water is one more victim of our present drought conditions.  During the winter, we often see water splashing over the weir at the outfall; today, the weir head is a metre above the lake level and flow in the River Ehen is maintained only by pumps installed by United Utilities.

When W.H. Pearsall visited Ennerdale Water in the 1920s, he considered it to be one of the most primitive of the Lake District’s lakes (see “The power of rock …”).   However, this supposedly wild lake had been tamed by a weir since the middle of the 19th century in order to provide drinking water for Whitehaven and Workington and surrounding areas.   That, in turn, has consequences for the river downstream, especially at times such as this when, unless augmented by pumps, there would be no water in the River Ehen below the outfall.   At some point in the next decade, a new water infrastructure project will pipe water to west Cumbria from Thirlmere, after which the weir can be removed and fluctuations in water level in both lake and river will be more natural.

The weir at the outfall of Ennerdale Water, with the fish pass at the far end. 

Meanwhile, however, I was able to explore areas of the lake littoral zone that would normally be hidden from me.  My notebook, for example, records my observations that this part of the lake shore has a stony bottom yet, as can be seen from the picture above, these form a belt about 20 metres wide, after which there is firm sand.   Normally, this would be close to the limit of safe wading but, today, I could walk out with just a pair of thin neoprene shoes.   Looking down, I could see a number of tufts of the alga Nitella flexilis growing in this sand.   I’ve written about this species before (see “Finding the missing link in plant evolution …”) and have seen it in the lake before, but not in this particular location. Standing with the lake water lapping against my shins I could bend down and take some photographs of these with my underwater camera that give this usually chilly location a semi-tropical feel.

It is a useful reminder to those of us who dabble in lake littoral zones and think that we understand their ecology that a lot happens beyond the depth in which we can safely wade.   Marco Cantonati and colleagues, for example, have found big changes in the composition of the algal flora of Alpine lakes when they used Scuba diving to explore the depths of their littoral zones.  No doubt, we would see similar changes if we were to try the same in the Lake District.


Cantonati, M., Scola, S., Angeli, N., Guella, G. & Frassanito, R. (2009).  Environmental controls of epilithic diatom depth-distribution in an oligotrophic lake characterized by marked water-level fluctuations.   European Journal of Phycology 44: 15-29.

Cantonati, M. & Lowe, R.L. (2014).  Lake benthic algae: toward an understanding of their ecology.  Freshwater Science 33: 475-486.


That’s funny …

The most exciting phrase to hear in science, the one that heralds new discoveries, is not “Eureka!” but “That’s funny”
Attributed to Issac Asimov

I have visited Croasdale Beck, in western Cumbria, twenty-eight times since 2015 and I thought I was beginning to understand it’s character (see “A tale of two diatoms” and “What a difference a storm makes”).   It is the unruly sibling of the River Ehen which, usually, offers a far less amenable environment for freshwater algae.  Last week, however, as we walked down the track towards the stream, we were confronted with the unexpected sight of a river bed that was bright green.  Our measurements, too, showed that not only was there a lot of algae in absolute terms, but there was far more here than we had measured in the River Ehen.  Usually, the situation is reversed, with the Ehen having more than Croasdale Beck.

Croasdale Beck at NY 087 170 looking upstream in April 2018.   The position of the gravel bar has shifted over the time that we have visited, with the wetted channel originally being at the right hand side, rather than being split into two.

It was hard to capture the extent of the algae growing on the river bed in a photograph, but the macroscopic image below captures the colour of the growths well, and you’ll have to use your imagination to scale this up to cover half of the stream bed.  Under the microscope, these growths turned out to be virtual monocultures of the green alga Draparnaldia glomerata.  This is common in clean rivers in spring time, and I often find it in the nearby River Ehen (see “The River Ehen in February”).  What my images do not show is the mucilage that surrounds the filaments.   In some cases, the growths can be almost jelly-like, so prolific is this mucilage.   One of the roles of this mucilage plays is to serve a matrix within which enzymes released by the fine hairs at the end of the filaments can act to release nutrients bound into tiny organic particles (see “A day out in Weardale …”).

Growths of Draparnaldia glomerata in Croasdale Beck (NY 087 170) in April 2018.  The upper image shows the filaments growing on submerged stones and the lower image shows the bushy side-branches growing from a central filament.  Scale bar: 100 micrometres (= 1/10th of a millimetre).

We also sample a site a couple of kilometres downstream on Croasdale Beck and, here again, the river bed was smothered in green growths.  I assumed that this, too, was Draparnaldia glomerata but, when I examined the filaments under the microscope, it turned out to be a different alga altogether: Ulothrix zonata (see “Bollihope Bhavacakra” and links therein).   There is little difference between the two sites that might explain this: the latter is slightly lower and is surrounded by rough pasture whilst the other is closer to the fells.   However, I have seen both Ulothrix zonata and Draparnaldia glomerata at several other sites in the vicinity, and a simplistic interpretation based on agricultural enrichment does not really work.

There were also a few obvious differences in the diatoms that I saw in the two samples.   In both cases, we sampled stones lacking green algae but, instead, having a thick brown biofilm.  Several taxa were common to both sites – Odontidium mesodon, for example (broadly confirming the hypothesis in “A tale of two diatoms …”) and Meridion circulare was conspicuous in both.   However, the lower site had many more cells of “Ulnaria ulna” than the upper site.   Again, there is no ready explanation but, at the same time, neither green algae or diatoms at either site suggests anything malign.

Filaments of Ulothrix zonata at Croasdale Beck (NY 072 161).   The upper filament is in a healthy vegetative state (although the cell walls are not as thickened as in many populations).  The lower filament is producing zoospores.   Scale bar: 25 micrometres (= 1/40th of a millimetre).

Diatoms in Croasdale Beck, April 2018.   a. upper site: note the abundance of Odontidium mesodon, plus cells of Gomphonema cf exilissimum, Achnanthidium minutissimum and Meridion circulare; b. lower site: note the presence of “Ulnaria ulna” as well as several of the taxa found at the upper site.   Scale bar: 25 micrometres (= 1/40th of a millimetre).  

So where does this take us?  I talked about the benefits of repeat visits to the same site in “A brief history of time wasting …” and I think that these data from Croasdale are making a similar point.  By necessity, most formal assessments of the state of ecology are based on very limited data, from which, at best, we get an estimate of the “average” condition of a water body over a period of time.  Repeat visits might lead to a more precise assessment of the “average” state but also give us a better idea of the whole range of conditions that might be encountered.  Here, I suspect, we chanced upon one of the extremes of the distribution of conditions.   Cold, wet weather in early spring delayed the growth of many plants – aquatic and terrestrial – as well as the invertebrates that graze them.   Then the period of warm, dry conditions that preceded our visit gave the algae an opportunity to thrive whilst their grazers are still playing “catch-up”.  I suspect that next time we visit Croasdale Beck will have its familiar appearance.   It is, nonetheless, sobering to think that this single visit could have formed fifty-percent of the evidence on which a formal assessment might have been made.


Small details in the big picture …

I’ve written about Platessa oblongella, a small diatom common in low alkalinity environments, before (see “A tale of two diatoms …” and links therein) but my travels around west Cumbria are gradually revealing more and more about the ecology of this organism, so bear with me as I explain my latest findings.

My first graph shows how the distribution of this diatom varies in different types of water body in the Ennerdale catchment.   I have analysed 223 samples from this small area over the past few years and, within this dataset, there is a very clear distinction between situations where Platessa oblongella is abundant and situations where it is very rare.   I have very few records from Ennerdale Water itself (present in just two out of 27 samples, and never comprising more than 2.7% of all diatoms in the sample) nor from the River Ehen, which flows out of the lake (present in just 16 out of 164 samples, and always £ 1% of all diatoms).  By contrast, in Croasdale Beck and in streams that flow into the north-west corner of the lake, it is present in 28 out of 32 samples, with a maximum relative abundance of 69%.   In ten samples it forms more than 10% of all diatoms present.   Several of my samples from the small streams were collected from just a few metres above the point where they joined the lake, which makes the distinction between these streams and the lake that much more intriguing.

My theory – based on data I showed in A  tale of two diatoms  is that Platessa oblongella is a species of disturbed habitats and that the littoral zone of a lake, whilst subject to some turbulence, is less disturbed than the rough world of an unregulated stream.  The contrast between the River Ehen immediately below the dam at the outfall of the lake and the various small tributary streams also supports this idea.

Differences in percentage of Platessa oblongella (including P. saxonica) in epilithic samples from Ennerdale Water and associated streams.  Data collected between 2012 and 2018 (along with one sample from River Ehen collected in 1997).   The photograph at the top of the post shows Ennerdale Water, photographed in January 2018.

Some of the populations I looked at seemed to consist of two distinct forms, one broader than the other.   This variability is quite common in Platessa oblongella and Carlos Wetzel and colleagues recently published a paper which suggests that these are, in fact, two distinct species.   When I first started looking at diatoms, John Carter, my mentor, used the name Achnanthes saxonica, but Krammer and Lange-Bertalot, in the revised Süsswassserflora, regarded this as a synonym of Achnanthes oblongella, a species first found in Thailand.   Wetzel’s study shows, as well as the difference in valve width, differences in the fine details of the striae between the two species.   They also decided that both species belonged in the genus Platessa, rather than Achnanthes.

Platessa oblongella (top) and P. saxonica (bottom) from Croasdale Beck, October 2017.  Scale bar: 10 micrometres (= 1/100th of a millimetre).

Valve width is, however, a very useful criterion, as the histograms below show.   The left hand graph shows a distinctly bimodal distribution of widths in specimens from Croasdale Beck, whilst the right hand graph shows a much tighter, and clearly unimodal, range.   This comes from another tributary stream flowing into the Ehen about 500 metres below the lake itself.  Quite why two species can co-exist in one stream but only one is present in another is not clear.

The modes of these populations are very close to the median widths for P. saxonica (narrow, ± 4/5 – 5 mm) and P. oblongella (broader, ± 6.5 mm) respectively but, as the left hand histogram shows, there is some overlap.    You might have trouble, for example, deciding whether a valve that was 5.5 mm wide was a “fat” P. saxonica or a “thin” P. oblongella.   My standard advice in situations such as this is that we should identify populations not individuals although, in the case of Croasdale Beck, this will still leave a grey area between the “fat” and “thin” valves where a judgement call is necessary.   In this case I think I could have done it because the P. saxonica valves in this stream tended to have a greater length:breadth ratio than those of P. oblongella, though I have not actually quantified this.

Width of valves in populations of “Achnanthes oblongella” from a) Croasdale Beck, and b) an unnamed tributary stream of the River Ehen, October 2017. 

There is more to say about the ecology of these species, but I have probably written enough for now.  I will leave you, for now, to bask in the rare sensation that occurs when diatom taxonomists make a situation clearer rather than more opaque, and return to this subject in a future post.


Carter, J.R. (1970).   Observations of some British forms of Achnanthes saxonica Krasske.  Microscopy: Journal of the Quekett Microscopical Club 31: 313-316.

Wetzel, C.E., Lange-Bertalot, H. & Ector, L. (2017).  Type analysis of Achnanthes oblongella Østup and resurrection of Achnanthes saxonica Krasske (Bacillariophyta).  Nova Hedwigia, Beiheft 146: 209-227.

The underwater world of Ennerdale Water …

I’ve tried to capture the world of microscopic benthic algae many times but never, until now, attempted the same effect with plankton.   The picture below illustrates the problem that I face: whereas the benthic flora are organised with, for the most part, a clear three-dimensional structure and known dependencies amongst organisms (species A, for example, being epiphytic on species B), plankton are randomly distributed in a very dilute solution.   My picture  below, which is based on four phytoplankton samples collected by the Environment Agency in the summers of 2014 and 2016.

A representation of the phytoplankton of Ennerdale Water with cells of Rhodomonas and Kephyrion depicted at a realistic density (c. 1000 – 2000 cells per millilitre).

I had to address two issues in producing this image, which is based on four phytoplankton samples collected by the Environment Agency in the summers of 2014 and 2016: depicting the phytoplankton cells at approximately the correct density and making sense of the list of names that appeared on the list.  Ennerdale Water is a very nutrient-poor lake and cell concentrations during the summer are in the order of 1000 to 2000 per millilitre.  That sounds a large number until you consider the scale at which we are working.   For simplicity, I assumed spherical cells of about 20 micrometres diameter (= 1/50th of a millimetre) at a density of 1000 cells/ml.    That equates to one cell per micrometre which is 1 mm x 1 mm x 1 mm.   Using these assumptions, each cell is 50 diameters distant from its nearest neighbour, which means the foreground of a picture should contain only two small cells and a lot of blue paint.

Next, I need to know what algae to paint and the problem here is that 85 per cent of the cells in the Environment Agency phytoplankton analyses were described as “picoplankton < 2 micrometres diameter” or “nanoplankton 2-20 micrometres diameter” (the latter divided into flagellates and non-flagellates).  There are, apparently, big difficulties in naming many of the cells found as preservation with Lugol’s Iodine coupled with the long time in storage before analysis can lead to loss of useful diagnostic features.   Cells in the nanoplankton category can, in theory, belong to any one of a number of groups of algae but If I focussed just on those organisms that could be named, I see that the Cryptophyta Rhodomonas lacustris var nannoplanctica (formerly R. minuta var. nannoplanctica) predominates, followed by Chrysophytes, of which Kephyrion is the most abundant.   So these are the two cells that I have put in the foreground.

I subsequently turned up a paper from 1912 by the father and son team of William and George West who looked at the phytoplankton of Ennerdale Water and a number of other lakes in the Lake District and Scotland.  The range of taxa that they found was quite different to that recorded in these recent surveys with samples dominated by desmids and almost no Chrysophytes or Cryptophytes recorded at all. That may, in part, be due to differences in methods – they collected samples using a “silken tow net”, which would probably have missed the very small Chrysophyta and Cryptophyta (an earlier paper by them tells us of the size of the nets but not the mesh itself) .  Some desmids that they found were found in the recent surveys but in much smaller quantities and it is possible that this was partly an artefact of the differences in sampling technique.  The idea of comparing count data from old papers with modern records is appealing but, in most cases, separating genuine changes in composition from differences introduced by sampling and analytical methods is always difficult.

Excuse these ramblings … there is, as you can see, not a lot of pictorial interest in the underwater world of an oligotrophic lake.   If you want excitement, tune into Blue Planet II, David Attenborough’s latest series for the BBC You will find sex and violence galore there.  The underwater world of Ennerdale Water is a quieter, more serene and certainly less televisual place.  Maybe that’s not such a bad thing …


Lund, J.W.G. (1948) A rarely recorded but very common British alga, Rhodomonas minuta Skuja. British Phycological Bulletin, 2:3, 133-139.

West, W. & West, G.S. (1909). The British freshwater phytoplankton, with special reference to the desmid-plankton and the distribution of British desmids.   Proceedings of the Royal Society of London Series B 81: 165-206.

West, W. & West, G.S. (1912).  On the periodicity of the phytoplankton of some British lakes.  Journal of the Linnaean Society, Botany 40: 395-432.

What a difference a storm makes …

I was back at Croasdale Beck last week and noticed a rather dramatic change to the meander just upstream from our regular sampling spot.   If you look at the photograph that heads the post “A tale of two diatoms …”, you’ll see the stream flowing around this meander.  Now, however, it has cut a new, shorter channel that bypasses the meander altogether.   We visited the stream just a few days after Storm Ophelia had passed through although, judging by the grass growing on the gravel of the abandoned meander, it was not necessarily this particular event that reshaped the stream.

Croasdale Beck is an unruly tributary of the River Ehen, rising on the fells above Ennerdale Water and tumbling down across rough grazing land and some semi-improved pasture (as in the picture above) before joining the Ehen in Ennerdale Bridge.   This is not the first time that we have seen conspicuous changes in the channel after a storm.  The magnitude of the flood is illustrated by the hydrograph below, which went off-scale for a period, as the discharge exceeded 3000 mega litres per day (300 MLD is the approximate limit for safe wading, in my experience).   I noticed that there was much less green algae present than we usually record at this time of year, although the diatom film was still quite thick.   Some of the stones that I picked up to sample had the slimy biofilm on the underside, suggesting that they had been recently rolled by the flooded river.   Croasdale Beck has no lake to buffer the rise and fall of the floodwaters and a huge amount of energy is carried down in a short period of time as the water surges downstream.

By the time we had arrived, the floodwaters had subsided and the sheep were contentedly grazing the surrounding land.  The stream itself was almost back to base flow (in contrast to the River Ehen which was still only just wadable).  Only the meander looked different …

The hydrograph for the River Ehen, as the aftereffects of Storm Ophelia make their way downstream.

Buffers for duffers …

In Ecology in the hard rock café I wrote about the challenges of living in an aquatic world where carbon – one of the raw materials for photosynthesis – was in short supply.   What I did not write about in that post is that this carbon also gives freshwater some useful additional properties.   In brief, rainwater is not pure water, but absorbs carbon dioxide from the atmosphere.  This, in turn, makes rainwater slightly acidic and, when it falls onto rocks, this weak acid dissolves the minerals from which the rock is made.  This adds two other forms of carbon to the water – bicarbonate and carbonate (the latter, particularly, from limestone).

Each of these three types of carbon in freshwater can convert to either of the other two types, with the speed of the reaction depending on the balance between the forms (the “law of mass actions”).  In essence, the reactions proceed until equilibrium is obtained, and this equilibrium, in turn, depends upon the pH of the solution.  These processes are summarised in the diagram below.

Relationship between pH and the proportion of inorganic carbon as free carbon dioxide (or carbonic acid, H2CO3 – orange line), bicarbonate (HCO3 – green line) and carbonate (CO32- – blue line).

The chemistry behind this is not easy to explain but a consequence is that any attempt to shift the pH (e.g. by adding acid) causes an automatic adjustment in the balance between the different forms of carbon.  Some of the hydrogen ions that could make the water acid are, instead , bound up as bicarbonate, and the pH, as a result, does not change.  The greater the quantity of inorganic carbon in the sample, in other words, the greater the capacity of the water to resist changes in pH.   The carbonate, bicarbonate and free carbon dioxide together act as a “buffer”, a chemical shock absorber.   Think of it as equivalent to the responsible use of a credit card or savings account to defer the cost of an unexpected bill (a car repair, for example) so that your current account does not go overdrawn.

Because life largely evolved in well-buffered marine systems, the enzymes that run our cells generally work best within a narrow range of pH (approximately 6-9).   Cells – unicellular life forms in particular – get stressed if pH strays outside this range, so the greater the buffering capacity, the easier it is for cells (life at high pH can bring additional complications, but we don’t have time to go into those here).  “Alkalinity”, as I mentioned in the earlier post, is the measure that ecologists use to assess the strength of the buffer system in a lake or river.  The principle of the measurement is straightforward: we add a dilute acid very slowly and watch what happens to the pH.   At first, nothing happens but, as soon as the water’s natural buffering capacity has been exceeded, pH drops rapidly.

I have a small portable alkalinity titration kit which involves adding drops of bromophenol blue indicator to a sample of stream or lake water.  This gives the water a blue colour when the pH is greater than 4.6.  As the pH falls, the solution becomes colourless and, eventually, turns yellow.   If you look at the graph above you will see that, at pH 4.6 most of the bicarbonate (HCO3) has been converted to carbon dioxide so the buffering capacity is pretty much non-existent.  This means that I can use the quantity of acid that is needed to make the bromophenol blue change colour as a measure of the buffering capacity of the water.

Alkalinity titrations beside Ennerdale Water (see top photograph) using a Hanna HI 3811 alkalinity test kit.  The right hand image shows acid being added to the water sample with a 1 ml pipette.  The blue colour shows that pH has not yet dropped below 4.6.

All this talk of chemical equilibria seems to be a long way from the natural history that is the core business of this blog.  Yet, at the same time, these reactions describe natural phenomena every bit as real as the plants and animals that attract the interest of naturalists.   Geology and chemistry ultimately create the context within which biology flourishes, but it is rare to meet a chemist who can talk with a naturalist’s passion.  I think that this is partly because chemistry tends not to describe tangible features of the landscape but, instead, quickly gets lost in abstract equations.  However, it is also a matter of culture: chemists need clinical separation from the mud and filth to maximise precision, whilst ecologists feel the lure of the field.  There is, nonetheless, a very basic and necessary link between the chemistry and ecology of aquatic systems.   Geology may shape a landscape but chemistry is one of the key mediators that determines the types of plants that cloak the hills and vales.  We ignore it at our peril.