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.

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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.

Desmid masterclass …

My chance encounter with desmids in the Pirin Mountains (described in the previous post) was a serendipitous insofar as, less than a fortnight after my return from Bulgaria I find myself standing beside a soft water tarn in the Lake District noted for its rich desmid flora.  I’m here for a joint field meeting of the British Phycological Society and Quekett Microscopical Club and, having decided on a venue, realised that the location of the Freshwater Biological Association’s laboratory meant that desmids were the obvious focus of our investigations.

So, on Friday afternoon, a small convoy of vehicles made its way from the FBA, on the west side of Windermere, via Hawkshead and Coniston, to two small tarns for an initial collecting trip.  Dave John, our in-house desmid expert, had been directed to these by David Williamson, the UK’s leading expert on desmids (but, alas, too elderly and infirm to join us), who had recorded large numbers of desmids from both Kelly Hall Tarn and Long Moss Tarn.   Fortunately, the weather today was rather more amenable to sampling than my previous desmid-hunting trip in this part of the world (see “Damp days in search of desmids”).

The predilection of desmids for boggy areas means that there is always a risk of damp feet when –collecting samples, especially when grabbing a chance sample whilst out walking.   Last time, my problem was that footwear appropriate to getting to Dock Tarn was not really appropriate for getting into Dock Tarn; today, however, I could tramp the short distance from our vehicles in wellies.  Dave noted wryly that most of David Williamson’s best sites were close to car parks, and that many of the sites sampled by the Wests, writers of a significant desmid flora of Britain in the late 19th century, were similarly placed closed to train stations.   Such practicalities probably lie behind more of our decisions about where to sample than we care to admit.

Once by the edge of the Kelly Hall Tarn, Dave gave us a demonstration of sampling desmids, starting with David Williamson’s preferred method using a plankton net.   There are two options – either casting a net attached to a piece of rope, then drawing it back through the water  or using a small plankton net attached to a telescopic handle.   Dave’s has a mesh of 33 micrometres (1/30th of a millimetre) which is fine for relatively large algae such as desmids.   He took care to avoid disturbing the bottom, in order to collect only phytoplankton but also pulled the through patches of submerged plants in order to dislodge the algae that live around their stems and leaves.  With care, it is also possible to brush the net gently against the bottom, dislodging some of the heavier desmids that typically sit just on the surface of the bottom sediments.  Too much disturbance will mean that the net quickly clogs with fine sediment particles.

Dave John demonstrating how to sample desmids with a plankton net at Kelly Hall Tarn, Cumbria, September 2017.

From Kelly Hall Tarn it was a short uphill walk to Long Moss Tarn for our second demonstration.   There is usually a part of the perimeter of tarns in this part of the world that is marshy, often dominated by Sphagnum.  The hummocks of Sphagnum were of no great interest except as stepping stones to the edge but, once there, we could see the semi-aquatic Sphagnum cuspidatum living at the margins of the lake (moss cognoscenti describe S. cuspidatum as looking like “drowned kittens”).   Dave took a handful of this soft, damp moss, flicked it a couple of times to remove surplus water, then gently squeezed it, catching the water that oozed out of the moss in a small sampling bottle.   Holding this up to the light, we could see a distinct green tinge which brought a big smile to Dave’s face, anticipating the rich haul of desmids that awaited us.

Dave John demonstrating how to collect desmids by squeezing Sphagnum at Long Moss Tarn, Cumbria, September 2017.

A patch of Sphagnum cuspidatum at Kelly Hall Tarn: perfect desmid habitat.

Kelly Hall Tarn had one further attraction for me.   It is perched on a hillside overlooking Coniston Water and from a small knoll a couple of hundred metres from the tarn I had a wonderful view southwards down the lake.  In the middle distance I could see Peel Island which was the inspiration for “Wild Cat Island” in Arthur Ransome’s Swallows and Amazons, a book that I read and re-read many times as a child.   At the north end of the lake you will find Bank Ground Farm, which was the setting for “Holly Howe” in the book and which, I notice, still offers accommodation and can be booked via Airbnb.   Finally, on this theme, the Steamboat Museum in Windermere has the original “Amazon”, the sailing dinghy that is the centre of many of the adventures.

Enough of these digressions.   We now have samples from two tarns and the next post will start to explore the contents of these and other samples that we collected.

The view down Coniston Water from Long Moss Tarn.  Peel Island is in the centre of the picture.

 

 

Damp days in search of desmids …

Seatoller, in Borrowdale, is the wettest place in England, so we should not have been surprised by the persistent drizzle that accompanied us as we set off hunting for desmids last week.  The combination of Borrowdale’s hard volcanic rocks and a damp climate combine to create ideal habitats for bog-loving desmids and I had intelligence that Dock Tarn, on the fells above Borrowdale, was a hot spot of desmid diversity.   Getting there, however, was no easy task.  Though just a couple of kilometres from Stonethwaite on the map, there were an awful lot of contour lines awfully close together between the beginning and end of our walk.   The footpath zig-zagged through ancient woodland clinging to a steep hillside until we emerged onto the moorland above.  We then made our way across a plateau covered with heather moorland until we saw the tarn stretching away into the mist in front of us.

You know you are in good desmid habitat when there is water percolating into your body from both ends: rain dripping down from the hood of your cagoule and dampness seeping in through your shoes.  They are organisms that love marshy, boggy conditions, especially in areas where the water is as soft as it is here.   The alternative to damp feet would be to either climb up from Borrowdale in Wellingtons or waders or carry them up that steep hillside in a rucksack.   However, I suspect that the mud at the bottom of the tarn was too soft and deep for Wellington boots and lugging waders up that hillside would have been hard work so damp feet was the price I had to pay.   I leaned out as far as I could from the shore to grab some of the sedge stems which had a visible coating of attached algae, and also squeezed the peaty water from a few handfuls of Sphagnum that I pulled from a boggy pool.  That would have to do on this particular morning as the rain was now soaking through my trousers and, in any case, there were places I needed to be later that morning.   I shoved the bottles containing my samples into my rucksack and followed the path back down the hillside.

Epiphytic algae growing around a sedge stem in the outflow of Dock Tarn, Cumbria, July 2017.   The width of the stem plus epiphytes is about half a centimetre.

Dock Tarn is one of a number of sites identified as an “Important Plant Area” (IPA) on the basis of the rich desmid flora, largely due to work over the years by David Williamson.   It qualifies as an IPA on four criteria: the presence of threatened species, high diversity, a long history of study and because it represents a “threatened habitat”.   David Williamson has recorded over 50 species from this location, 13 of which are candidates for a “potential Red Data List”.   A few of these are illustrated in the figures below.   One of the species in the first image, Haplotaenium minutum, belongs to a genus only recently separated from Pleurotaenium, which looks very similar to the untrained eye (the difference lies in the structure of the ridges on the chloroplast).  Looking at these long cylindrical cells serves to emphasise just how much dexterity Chris Carter needed to produce his Hilda Canter-Lund prize winning image.  Images in the second plate include two more species of the genus Xanthidium, which we met in “Desmids on the defensive …”.

Dock tarn desmids: a. Netrium digitus var. latum; b. Tetmemorus brebissonii; c. Haplotaenium minutum.  Scale bar: 25 micrometres ( = 1/40th of a millimetre). 

The desmids in the lower plate, in particular, show one of their key characteristics very clearly: their cells are divided into two distinct lobes (“semicells”) joined by an isthmus (the word desmid comes from the Greek desmos, meaning “bond”).  The image of Staurastrum manfeldtii var. productum also shows a number of bacteria growing on the cell: these are probably growing within the mucilage that desmids secrete around themselves whilst there are distinct pyrenoids in the two Xanthidium species.  Their predilection for soft water means that they need the carbon-concentrating mechanisms that these contain if they are to thrive.   Not all desmids live in water as soft as this, and some are able to use inorganic bicarbonate to fuel their photosynthetic engine, but there will be little or no bicarbonatae in a habitat such as Dock Tarn.   I wrote about these carbon concentrating mechanisms in algae from Ennerdale Water (see “Concentrating on carbon …”) and the two filamentous algae that featured in that post, Mougeotia and Spirogyra, both belong to the same class within the green algae as the desmids (Conjugatophyceae or Zygnemtetophyceae).

There will be more about desmids on this blog over the next few months in preparation for a the weekend of 15-17 September when I am organising a joint meeting of the British Phycological Society and Quekett Microscopical Club in Windermere.  We’ll be visiting some other Lake District tarns known to be rich in desmids during this weekend and have Dave Johns and Allan Pentecost on hand, amongst others, to offer expert advice on what we find.  There are still a few places left, so hurry up to book your place.  I haven’t done a great job of selling the Cumbrian climate in this post but we have the use of the Freshwater Biological Association facilities, including a laboratory and the library, so no one need get damper than they want.   See you there…

More desmids from Dock Tarn: d. Euastrum cuneatum; e. Xanthidium cristatum var. uncinatum; f. Xanthidium antilopaeum; g. Staurastrum manfeldtii var. productum.   Scale bar: 25 micrometres
( = 1/40th of a millimetre). 

References

Coesel, P.F.M. (1994). On the ecological significance of a cellular mucilaginous envelope in planktic desmids. Algological Studies 73: 65-74.

Kiemle, S.N., Domozych, D.S. & Gretz, M.R. (2007). The extracellular polymeric substances of desmids (Conjugatophyceae, Streptophyta): chemistry, structural analyses and implications in wetland biofilms. Phycologia 46: 617-627.

Spijkerman, E., Maberly, S.C. & Coesel, P.F.M. (2005).  Carbon acquisition mechanisms by planktonicdesmids and their link to ecological distribution. Canadian Journal of Botany 83: 850–858.

 

The exception that proves the rule …

If you are going to understand river ecology, you need to be able to consider landscapes at several different scales simultaneously.   In the River Ehen, this means looking upstream towards Ennerdale Water and, beyond, to Great Gable and the other Lake District peaks in order to appreciate the geology that gives the catchment its bones.  But, at the same time, you need to look around at the meanders of the river and the bankside vegetation that create the immediate habitat for the organisms, and then to look even more closely at the individual stones that line the river bed.

Peering into the water last week, the pebbles, cobbles and boulders that make up the substratum of the River Ehen looked bare of filamentous algae for the most part.  There were a few clumps but, at this time of year, when grazing invertebrates are active, the algal flora is reduced to a thin film, invisible to the naked eye and apparent only as a slimy sensation when you run your fingers across the stone’s surface.   However, when I picked up a couple of cobbles, I noticed small, pale green gelatinous growths stuck on the upper surface.   Most were just a few millimetres across with the largest up to about a centimetre.

A growth of Draparnaldia glomerata on the upper surface of a cobble in the River Ehen, Cumbria, April 2017.

These growths are composed of the green alga Draparnaldia glomerata.  I have written about this alga before (see “The River Ehen in February”) but, under the microscope, it is such a beautiful organism, that I am not going to apologise for writing about it again.   The alga lives inside the gelatinous mass and consists of a relatively thick central filament from which tufts of narrower side-branches emerge.  The cells that make up these side branches gradually narrow, and the chloroplast becomes smaller until, eventually, the cells form a colourless “hair”.   These hairs are relatively short on the material illustrated below but can be much longer (some longer hairs were present but did not present nicely for photography).  The hairs are, in fact, an adaptation to help the alga acquire phosphorus, something I described in an earlier post about a relative, Stigeoclonium tenue (see “A day out in Weardale”).

Draparnaldia glomerata from the River Ehen, April 2017 showing filaments and side branches. Scale bars: a.: 50 micrometres (= 1/20th of a millimetre); b.: 20 micrometres (= 1/50th of a millimetre).

A low concentration of phosphorus is usually regarded as a Good Thing by aquatic ecologists, as this limits the amount of energy produced  by the plants at the base of the food chain.  This, in turn, means that the microbes and animals that depend on these are not using up all the oxygen in the water, or having other deleterious influences on the ecosystem.   I would usually regard the presence of an organism such as Draparnaldia as a sign of a healthy stream, as it is adapted to thrive when phosphorus is relatively scarce.

I was, however, careful to place “relatively” in front of “scarce”.   Studies by my colleagues (referenced in the earlier post) showed that the production of the phosphatase enzyme that boosts the alga’s ability to acquire phosphorus when it is scarce is determined by the ratio of nitrogen to phosphorus inside the cell itself, rather than in the water.   The physiology of nutrient limitation is all about the balance between the different “ingredients” that a cell needs.   If you have three eggs and 170g of sugar, for example, you can only make one cake, regardless of how much flour you have in your cupboard.   So it is with algae: most of the locations where I find Draparnaldia have very little nitrogen, but even less phosphorus.   There are barely enough ingredients for the algal “cake” so it is advantageous to the organism to pump out some enzyme to order to make up the shortfall.  This means that I can say with confidence that Draparnaldia is usually a good indicator of healthy streams.

Just occasionally, however, I get Draparnaldia in places where I would not usually expect it to be found.   The picture below shows a colleague standing in the Terman River, just before it flows into Lough Erne in Northern Ireland.   She is holding a skein of Cladophora glomerata in her left hand and a skein of Draparnaldia in her right hand.  I associate the former with nutrient-rich rivers where I would not usually expect to find Draparnaldia.  But both were growing prolifically at this site which defied my expectations until I started to think about the physiology of the organism.   Had I had the facilities to analyse the tissues of the algae, I expect that I would have found very high concentrations of nitrogen which, in turn, creates a demand for yet more phosphorus so that it could convert that nitrogen into the proteins it needs to grow.  However, that cannot be the whole story, because normally, under such circumstances, I would expect a competitive alga such as Cladophora to out-compete and overgrow the Draparnaldia.   Here, they were growing side-by-side.   It is, to date, the most luxuriant growth of Draparnaldia that I have seen, and also the only occasion where I have seen these two species co-existing in such abundance.

My colleague, Bernie White, holding skeins of Cladophora glomerata (left hand) and Draparnaldia glomerata (right hand) from the Terman River near Toome.  The border between the Republic of Ireland and the UK runs along the middle of this river.

I can extend my lesson from the first example to say that, to understand the ecology of any particular river you need to have perspectives obtained from many other rivers.   But, in this case, we see a potential limitation: the case of the “rare exception” that clouds an otherwise clear picture of an association between an organism and a particular set of circumstances.   The problem is particularly acute when dealing with the effect of nutrients because we are usually dealing with indirect, rather than direct effects.   Draparnaldia glomerata is usually associated with clean rivers with low concentrations of nutrients but it is not there because nutrient concentrations are low.   As for the diatom Amphora pediculus (see “The challenging ecology of a freshwater diatom?”) a more nuanced understanding of the relationship between an organism and nutrients yields more useful insights than simply assuming a cause-effect relationship.