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 …

References

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.

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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.

Winning hearts and minds …

I write several of my posts whilst travelling, though am always conscious of the hypocrisy of writing an environmentally-themed blog whilst, at the same time, chalking up an embarrassing carbon footprint.  Last month, however, I participated in my first “eConference”, in which the participants were linked by the internet.  With over 200 people from all over Europe, and beyond, attending for all or part of the three days, there was a substantial environmental benefit and whilst there was little potential for the often-useful “off-piste” conversations that are often as useful as the formal programme of a conference, there were some unexpected benefits.  I, for example, managed to get the ironing done whilst listening to Daniel Hering and Annette Battrup-Pedersen’s talks.

You can find the presentations by following this link: https://www.ceh.ac.uk/get-involved/events/future-water-management-europe-econference.   My talk is the first and, in it, I tried to lay out some of the strengths and weaknesses of the ways that we collect and use ecological data for managing lakes and rivers.  I was aiming to give a high level overview of the situation and, as I prepared, I found myself drawing, as I often seem to do, on medical and health-related metaphors.

At its simplest, ecological assessment involves looking at a habitat, collecting information about the types of communities that are present and match the information we collect to knowledge that we have obtained from outside sources (such as books and teachers) and from prior experience in order to guide decisions about future management of that habitat. At its simplest, this may involve categoric distinctions (“this section of a river is okay, but that one is not”) but we often find that finer distinctions are necessary, much as when a doctor asks a patient to articulate pain on a scale of one to ten.  The doctor-patient analogy is important, because the outcomes from ecological assessment almost always need to be communicated to people with far less technical understanding than the person who collected the information in the first place.

I’ve had more opportunity than I would have liked to ruminate on these analogies in recent years as my youngest son was diagnosed with Type I diabetes in 2014 (see “Why are ecologists so obsessed with monitoring?”).   One of the most impressive lessons for me was how the medical team at our local hospital managed to both stabilise his condition and teach him the rudiments of managing his blood sugar levels in less than a week.   He was a teenager with limited interest in science so the complexities of measuring and interpreting blood sugar levels had to be communicated in a very practical manner.  That he now lives a pretty normal life stands testament to their communication, as much to their medical, skills.

The situation with diabetes offers a useful parallel to environmental assessment: blood sugar concentrations are monitored and evaluated against thresholds.  If the concentration crosses these thresholds (too high or too low), then action is taken to either reduce or increase blood sugar (inject insulin or eat some sugar or carbohydrates, respectively).   Blood sugar concentrations change gradually over time and are measured on a continuous scale.  However, for practical purposes they can be reduced to a simple “Goldilocks” formula (“too much”, “just right”, “not enough”).  Behind each category lie, for a diabetic, powerful associations that reinforce the consequences of not taking action (if you have even seen a diabetic suffering a “hypo”, you’ll know what I mean).

Categorical distinctions versus continuous scales embody the tensions at the heart of contemporary ecological assessment: a decision to act or not act is categorical yet change in nature tends to be more gradual.   The science behind ecological assessment tends to favour continuous scales, whilst regulation needs thresholds.  This is, indeed, captured in the Water Framework Directive (WFD): there are 38 references to “ecological status”, eight in the main text and the remainder in the annexes.  By contrast, there are just two references to “ecological quality ratios” – the continuous scale on which ecological assessment is based – both of which are in an annex.   Yet, somehow, these EQRs dominate conversation at most scientific meetings where the WFD is on the agenda.

You might think that this is an issue of semantics.  For both diabetes and ecological assessment, we can simply divide a continuous measurement scale into categories so what is the problem?   For diabetes, I think that the associations between low blood sugar and unpleasant, even dangerous consequences are such that it is not a problem.  For ecological assessment, I’m not so sure.  Like diabetes, our methods are able to convey the message that changes are taking place.  Unlike diabetes, they are often failing to finish the sentence with “… and bad things will happen unless you do something”.   EQRs can facilitate geek-to-geek interactions but often fail to transmit the associations to non-technical audiences – managers and stakeholders – that make them sit up and take notice.

I’d like to think that we can build categorical “triggers” into methods that make more direct links with these “bad things”.  In part, this would address the intrinsic uncertainty in our continuous scales (see “Certainly uncertain …”) but it would also greatly increase the ability of these methods to communicate risks and consequences to non-technical audiences (“look – this river is full of sewage fungus / filamentous algae – we must do something!”).   That’s important because, whilst I think that the WFD is successful at setting out principles for sustainable management of water, it fails if considered only as a means for top-down regulation.   In fact, I suspect that Article 14, which deals with public participation, is partly responsible for regulators not taking action (because “costs” are perceived as disproportionate to “benefits”) than for driving through improvements.   We need to start thinking more about ensuring that ecologists are given the tools to communicate their concerns beyond a narrow circle of fellow specialists (see also “The democratisation of stream ecology?”).   Despite all the research that the WFD has spawned, there has been a conspicuous failure to change “hearts and minds”.  In the final analysis, that is going to trump ecological nuance in determining the scale of environmental improvement we should expect.

It’s all about the algae

Just a short post to point you all towards an article I wrote for Royal Society of Biology’s magazine The Biologist.  It is a broad overview of the reasons why we use algae to assess the condition of our lakes and rivers in Europe and is illustrated with three of Chris Carter’s beautiful images, and the print edition will have even more of these.  Take the figure legends with a pinch of salt (we didn’t write these!): neither Tolypella nor Chaetophora are particularly common in the UK.   Navicula, on the other hand, is common but the legend makes no mention of this.

Whilst I have your attention, I will also point you towards a short article that I wrote for the most recent Phycological Bulletin, the newsletter of the Phycological Society of America.  This offers a few more hints to anyone thinking about entering the Hilda Canter-Lund competition next year.

Taking desmids to the next dimension …

Participants at the British Phycological Society / Quekett Microscopical Club field weekend at the Freshwater Biological Association, September 2017.  Scale bar: one metre (= 1,000,000 micrometres).

A theme that has run through this blog over the years has been that what you see down a microscope is often a highly distorted view of reality and at the end of our weekend of desmid hunting, Chris Carter gave a talk that also made this point, using desmids as a case study.  In essence, we had spent much of Saturday and Sunday morning peering down microscopes at three-dimensional objects that appeared, as a result of the very shallow depth of field that is characteristic of high magnification images, two dimensional.   We were then matching these to two-dimensional representations in the Floras and identification guides that we had to hand.  Dave explained a few tricks that experts use, such as applying gentle pressure to a coverslip with a fine needle, to turn desmids in order to see them from other angles but, mostly, we were restricted to very flattened views of desmids.

Chris has tackled this problem from several directions over the years, including experiments with anaglyphs (see “Phworrrrhhh …. algal sex in 3D!“) as well as the very careful manipulation of a long, cylindrical Pleurotaenium that won the Hilda Canter-Lund prize earlier this year.   He has also produced a number of plates with desmids laid out almost as if on an engineer’s drawing board, with front, top and side views.   Several of these are on Algaebase, but one example is reproduced below.  Microscopists learn to use the fine-focus control to appreciate the depth of the objects that they are examining and Chris also shows how it can reveal the nature of surface ornamentation on different parts of the cell.  The temptation, given a series of photos such as these (excluding the side view) would be to use “stacking” software to produce a single crisp image.  This is appropriate in some situations but you are, in truth, just producing a crisp two-dimensional image rather than offering any insights into the true shape of the cell.

Staurastrum furcatum from Botswana, photographed by Chris Carter.

Another technique that can be used to generate three-dimensional images is, of course, scanning electron microscopy.  However, this is beyond the budget for anyone outside a major institution.  This has helped greatly get a better understanding of the morphology of diatoms, in particular, but the third dimension comes at a price.   Scanning electron micrographs take us to an opaque, monochrome world, purged of the vivid colours that the microscopic world usually offers us.

Chris’ pièce de résistance, however, was a three-dimensional model of a Staurastrum, produced by the 3D printing company Shapeways and loosely-based on various pictures of S. furcatum and presented to him as a 70th birthday present by his son.  The main point is to demonstrate the symmetry and gross features of a typical Staurastrum rather than to be a taxonomic blueprint. The designers were very helpful but it does hint at what is possible with modern technology.

Chris Carter’s three-dimensional model of Staurastrum.  It is about six centimetres across.   You can buy your own copy from Shapeways by following this link …

The missing ingredient in this recipe is imagination.  Or, to be more precise, the viewer’s imagination as Chris has clearly demonstrated that he is not lacking in that department.  Once you have a sense of the three-dimensional form of a Staurastrum, you be able to use that knowledge every time you look at a two-dimensional image of a desmid through a microscope.   Seeing, as Ernest Gombrich reminds us in his great book Art and Illusion, is as much about using prior experiences to interpret the raw data collected by our optic nerves as it is about the patterns of light that stimulate our retinas.   Just as a child can look at a two-dimensional image of a cat in a book and match this to the real creatures that he or she encounters, so knowing about Staurastrum’s third dimension helps us to interpret the flat shapes that we see.

At a more basic level, all identification is a matter of matching the objects we see either to schemata stored in our memory or to patterns in books.  This, in turn, helps us to understand why the microscopic world seems so strange and mysterious to those who do not study it.  It all comes down to having (or not having) the prior experiences that generate recognition.   At one level, there are gasps of astonishment as people with none of these schemata in their memories encounter the beauty of desmids for the first time.  And then there is Frans Kouwets, another speaker at the meeting , who is busy cataloguing 750 different species of one genus, Cosmarium.   And in between there are the rest of us …

Frans Kouwets explains his fascination with Cosmarium to the British Phycological Society / Quekett Microscopical Club field meeting at the Freshwater Biological Association in September 2017.

Different tarn, different desmids …

Geoff and Chris, two of our band of desmid hunters, chose to stay in the FBA’s brand new holiday apartments and, rather than cross the lake to join us on Saturday morning they headed out to Moss Eccles Tarn, in the area between Esthwaite Water and Windermere.   One of Dave’s first dips into one of their samples yielded an almost pure monoculture of another filamentous desmid, Spherozosma vertebratum which presented some beguiling abstract patterns on my computer monitor.

Spherozosma vertebratum from Moss Eccles Tarn, September 2017.   Scale bar: 25 micrometres (= 1/40th of a millimetre).

Curiously, after our first encounter with Spherozosma vertebratum we did not see it in any of our other dips into the Moss Eccles samples although there were plenty of other desmids on display.   The most abundant of these was Staurastrum productum and, usefully, there were examples showing both apical and side views.   The three arms are distinctive (and distinguish it from relatives such as S. arachne which have five) and you can also see the knobbly “verrucae” on the spines as well as a broad mucilaginous envelope around the cells.

Staurastrum productum in side (left) and apical (right) views.  Images photographed from a computer monitor so apologies for their poor quality.  Scale bar: 25 micrometres (= 1/40th of a millimetre).

Another desmid with spines and mucilage was quite common.  This was Staurodesmus bulnheimii.  Spines slow the rate of sinking so are associated with several genera of predominately planktonic desmids.   The star-shaped arrangement of colonies of the diatom Asterionella formosa play a similar role (see “Little bugs have littler bugs upon their backs to bite ‘em”).   There were also several cells  of a small Cosmarium species, including some that had recently divided and the image shows how one cell has split down the central isthmus and a new semicell is growing back on each of the two daughter cells.   Finally, I have included an illustration of Micrasterias radiosa.  To the uninitiated this may look little different to M. compereana, illustrated in the previous post, but if you look closely you will see that the incisions between the lobes are much deeper in M. radiosa.

One sample from Moss Eccles Tarn kept me busy for half the morning and this account describes only part of the diversity.   Note how the differences between this and the School Knott Tarn sample are not just in the genera and species present but also in the life-forms I found.  The School Knott sample was from a Sphagnum squeezing whilst the Moss Eccles sample was from a plankton net.  That explains why I saw more spine-bearing desmids in the latter.  If I had looked at a plankton sample from School Knott and a Sphagnum squeezing from Moss Eccles, I might have found a different balance of life-forms between the two tarns.   But time was running out and I had to move on …

More desmids from Moss Eccles Tarn, September 2017: a. Staurodesmus bulnheimii; b. Cosmarium quadrifarium var. hexastichum; c. Euastrum cf. gemmatum.   Scale bar: 25 micrometres (= 1/40th of a millimetre).

Micrasterias radiosa from Moss Eccles Tarn, September 2017.   Scale bar: 25 micrometres (= 1/40th of a millimetre).

Lessons from School Knott Tarn …

As not everyone could join us on our excursion on Friday afternoon, we repeated the exercise on Saturday morning, heading to a small tarn just a short walk from Windermere and Bowness.   Despite its proximity to two of the busiest towns in the Lake District, there were very few other people around to disturb our peace whilst we collected samples.   As at Kelly Hall and Long Moss Tarns, Dave had his plankton net out, but we also explored a boggy region at one end, finding more patches of Sphagnum but also extensive growths of Utricularia minor (Lesser Bladderwort), one of a small number of aquatic carnivorous plants.   Dave was particularly pleased by this find as he associates this particular plant with rich hauls of desmids.

It was tempting to linger in the sunshine beside School Knott Tarn but the green tinge of the water that dripped out of the Sphagnum squeezings in particular was enough to lure us towards the Freshwater Biological Association’s laboratories in order to start examining our samples.

Utricularia minor (Lesser Bladderwort) from School Knott Tarn, near Windermere, September 2017.   Several of the spherical bladders which trap small invertebrates are visible on the plant.

My selection of photographs below shows just a part of the diversity that we encountered during our microscopic examinations.  I was using a borrowed set-up and the images are all from photographs of the desmids displayed on computer monitor, which is far from ideal.   Some of the larger desmids – one large Closterium species in particular – were too large to fit onto the screen and have had to be omitted from this account.  There were also a number of cells of Eremosphaera (see “More from Loughrigg Fell”) and some Cyanobacteria (Merismopedia was quite common) so this is a very partial description of our microscopical adventures in School Knott Tarn.

The first two desmids, Spirotaenia condensata and Cylindrocystis gracilis, belong to a group of desmids called “saccoderm desmids”.  These are more closely related to filamentous green algae of the Zygnemetaceae that are old friends of this blog (see “Concentrating on Carbon, for example) and, in fact, we could think of these genera as being unicellular analogues of their filamentous cousins.   Spirotaenia, with its helical chloroplast, for example, recalls Spirogyra whilst Cylindrocystis’ two star-shaped chloroplasts is reminiscent of Zygnema.  Mesotaenium, which we did not see in this sample, has a plate-like chloroplast similar to that in Mougeotia.

The next two illustrations both show species of Micrasterias.  Of these, M. compereana generated a vigorous discussion amongst our experts. This would have been described as M. fimbriata using the latest British floras but a paper has been published recently which uses molecular data to demonstrated the need to split the species. Finally, we have representatives of Euastrum and Haplotaenium, two genera that we also met at Dock Tarn (see “Damp days in search of desmids …”) although the species are different.   Haplotaenium differs from Pleurotaenium in the number and form of the chloroplasts and also because it lacks a terminal vacuole.

Desmids from Sphagnum squeezings from School Knott Tarn, September 2017: a. Spirotaenia condensata; b. Cylindrocystis gracilis; c. Micrasterias compereana; d. Micrasterias crux-meltensis; e. Euastrum oblongum; f. Haplotaenium rectum.  Scale bar: 25 micrometres (= 1/40th of a millimetre).

Four more desmids are illustrated on the lower plate.   Of these, we have seen Netrium digitus in Dock Tarn and the illustration there is better than this one, showing the undulating nature of the chloroplast margins quite clearly.   The desmid below this, Closterium closterioides caused some confusion at first.   We usually associate Closterium with lunate (moon-shaped) cells (see “More from Loughrigg Fell”) but this species is straight, sending me towards the section on Netrium in my Flora.  However, Netrium lacks terminal vacuoles whereas this specimen has prominent vacuoles at both ends.   We also found a variety, C. closterioides var. intermedium, in the same sample.

The final desmid that I have illustrated is a filamentous form: Desmidium schwartzii.  In contrast to Hyalotheca dissilens (see “Desmids from the Pirin mountains”) there is no obvious mucilaginous sheath around this specimen, but this may be an anomaly of this population or an artefact of the microscopy set-up.   We are looking at the side view of a chain of cells but if we were to look at the end view of one cell it would be triangular in this particular species.  The chloroplast fills most of the cell and has projections running into the corners of the cells.  However, as the filaments of the cells are slightly twisted, these projections appear to shift in position from cell to cell, giving a helical appearance.  I’ve tried to illustrate this with a schematic diagram.

More desmids from Sphagnum squeezings from School Knott Tarn, September 2017: g. Netrium digitus; h. Closterium closterioides var. closterioides; i. C. closterioides var. intermedium; j. Desmidium schwartzii Scale bar: 25 micrometres (= 1/40th of a millimetre).

This short post gives some idea of the diversity in a single sample from a single Tarn.   Dave handed all the samples we collected over to David Williamson on his way back south and we’ll get a fuller list of their diversity in due course.  This one sample occupied me for the latter part of Saturday morning and all of the afternoon.   On Sunday, I moved on to look at another sample and I’ll write about that in another post very soon.

A schematic view of a chain of Desmidium cells, showing the arrangement of the chloroplast seen in apical view (k.) and the implications of slight twisting of the filament on appearance (l.).  Diagram adapted from John et al. (2011).

Reference

John, D.M., Whitton, B.A. & Brock, A.J. (2011). The Freshwater Algal Flora of the British Isles. 2nd Edition. Cambridge University Press, Cambridge.

Neustupa, J., Šťastný, J. & Škaloud, P. (2014). Splitting of Micrasterias fimbriata (Desmidiales, Viridiplantae) into two monophyletic species and description of Micrasterias compereana sp. nov.  Plant Ecology and Evolution 147: 405-411.