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.

 

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Desmid diversity …

Back in September, I wrote about a joint British Phycological Society and Quekett Microscopical Club field weekend looking at desmids in the Lake District (see “Desmid Masterclass”, “Lessons from School Knott Tarn” and “Different tarn, different desmids …”).  Dave John sent some of the samples that we collected to David Williamson, the UK’s leading expert on desmids but, at 92, too frail to join us, and he has now sent back some fine drawings illustrating the range of desmids that he encountered.

Two of the tarns (Long Moss Tarn, Kelly Hall Tarns) are already recognised as Internationally Important Plant Areas (IPAs) for desmids because of their desmid diversity and containing internationally very rare desmids (based largely on David Williamson’s records) so their diversity is not a complete surprise to us.  Nonetheless, David found a total of 129 desmid taxa in the three tarns, whilst another desmid specialist, Marien van Westen, identified almost 160 desmids in another set of samples from the same tarns.

The drawings are arranged in three plates, one for each tarn.   Desmids identified by David Williamson from the three tarns are illustrated.  The desmids have been numbered and the captions prepared by David John who is analysing the findings and comparing them with surveys dating back to the 1970s.   David Williamson has drawn the taxa at different scales to roughly balance the arrangement on the collage, and adjusted the sizes so important details are visible.   No details of the chloroplasts are given since all samples had been preserved in formalin.  A few of the desmids, particularly those that are very long, have not been included in the plates.

Desmids from Long Moss Tarn (SD 292 936), September 2017.   Long Moss Tarn is shown in the photograph at the top of this post.

Desmids from Kelly Hall Tarn (SD 289 933), September 2017.

Desmids from School Knott Tarn (SD 427 973), September 2017.

Key

1-Actinotaenium diplosporum; 2-Actinotaenium turgidum;  3-Bambusina borreri;  4-Closterium acerosum var. borgei; 5-Closterium angustatum;  6-Closterium archerianum var. pseudocynthia;  7-Closterium archerianum; 8-Closterium attenuatum;  9-Closterium baillyanum var. alpinum; 10-Closterium baillyanum; 11-Closterium closterioides; 12-Closterium costatum; 13-Closterium dianae var. arcuatum; 14-Closterium dianae var. minus;  15-Closterium didymotocum; 16-Closterium incurvum; 17-Closterium intermedium; 18-Closterium kuetzingii;  19-Closterium lunula; 20-Closterium navicula;  21- Closterium setaceum; 22-Closterium striolatum; 23-Cosmarium amoenum; 24-Cosmarium anceps; 25-Cosmarium binum; 26-Cosmarium brebissonii; 27-Cosmarium contractum;  28-Cosmarium davidsonii; 29-Cosmarium debaryi;  30-Cosmarium depressum; 31-Cosmarium formosulum; 32-Cosmarium hostensiense; 33-Cosmarium incrassatum var. schmidlei; 34-Cosmarium margaritatum; 35-Cosmarium margaritiferum; 36-Cosmarium monomazum var. polymazum;  37-Cosmarium obtusatum;  38-Cosmarium ornatum; 39-Cosmarium ovale;  40-Cosmarium pachydermum; 41-Cosmarium pachydermum var. aethiopicum; 42-Cosmarium perforatum var. skujae; 43-Cosmarium portianum; 44-Cosmarium punctulatum;  45-Cosmarium quadratum; 46-Cosmarium quadrum; 47-Cosmarium subochthodes var. majus; 48-Cosmarium subtumidum var. groenbladii;  49-Cosmarium subundulatum; 50-Cosmarium tetragonum var. ornatum ; 51-Cosmarium tetraophthalmum; 52-Cosmarium variolatum;  53-Cosmocladium tuberculatum; 54-Desmidium aptogonum; 55-Desmidium swartzii; 56-Docidium baculum; 57-Euastrum ampullaceum; 58-Euastrum ansatum;  59-Euastrum bidentatum var. speciosum; 60-Euastrum gemmatum; 61-Euastrum luetkemulleri; 62-Euastrum oblongum; 63-Euastrum pectinatum; 64-Euastrum pulchellum; 65-Euastrum verrucosum; 66-Gonatozygon aculeatum; 67-Gonatozygon brebissonii; 68-Groenbladia undulata; 69-Haplotaenium minutum;  70-Hyalotheca dissiliens;  71- Micrasterias americana var. boldtii; 72-Micrasterias compereana; 73-Micrasterias crux-melitensis; 74-Micrasterias denticulata; 75-Micrasterias furcata; 76-Micrasterias pinnatifida;  77-Micrasterias radiosa; 78-Micrasterias rotata; 79-Micrasterias thomasiana; 80-Micrasterias truncata; 81-Netrium digitus; 82-Netrium digitus var. latum; 83-Netrium interruptum;  84-Penium exiguum; 85-Penium margaritaceum; 86-Pleurotaenium coronatum var. robustum;  87-Pleurotaenium ehrenbergii; 88-Pleurotaenium truncatum; 89-Sphaerozosma filiforme; 90-Staurastrum arachne;  91-Staurastrum arctiscon; 92-Staurastrum bieneanum; 93-Staurastrum boreale var. robustum; 94-Staurastrum cristatum; 95-Staurastrum dilatatum; 96-Staurastrum inconspicuum; 97-Staurastrum kouwetsii; 98-Staurastrum lapponicum; 99-Staurastrum maamense; 100-Staurastrum polytrichum; 101-Staurastrum productum; 102-Staurastrum quadrangulare; 103-Staurastrum striolatum; 104-Staurastrum teliferum; 105-Staurastrum tetracerum; 106-Staurodesmus convergens; 107-Staurodesmus convergens var. wollei; 108-Staurodesmus cuspidatus var. curvatus; 109-Staurodesmus megacanthus; 110- Xanthidium antilopaeum; 111-Xanthidium antilopaeum var. laeve; 112-Xanthidium antilopaeum var. polymazum; 113-Xanthidium cristatum.

More about Platessa oblongella and Odontidium mesodon

As my last post used the conventions of figurative art to describe algal ecology, I thought I would stick to graphs – science’s very own school of abstract art – for this one.   I spent some time in “Small details in the big picture” discussing the ecology of Platessa oblongella (including P. saxonica) but without saying very much about the types of streams where these species were found.  So I am going to take a step away from the Ennerdale catchment in this post and, instead, collate environmental data a large number of sites to get a broader understanding of their habitat preferences.  As these species are often associated with Odontidium mesodon (see “A tale of two diatoms …”), I will summarise the preferences of this species at the same time (but see Annex 1 for a graph of this species’ preferences for still versus standing water).

The first set of graphs show the response of these species to pH and alkalinity and establish both as species typical of circumneutral soft water.  Platessa oblongella can be abundant in more acid conditions (i.e. to the left of the green vertical lines) but most of the records where it is abundant have pH values between 6.5 and 7.5.   Note that P. oblongella can also be found in humic waters, where lower pH thresholds apply (see Annex 2).

Distribution of Odontidium mesodon and Platessa oblongella (including P. saxonica) to pH and alkalinity in UK streams.   Vertical lines for pH indicate threshold values that should support high (blue), good (green), moderate (orange) and poor (red) ecological status classes.  See Annex 2 for more explanation.

The second set of graphs shows how these species respond to inorganic nutrients.   Both are most abundant when inorganic nutrients are present in low concentrations, though the trend is stronger for phosphorus than it is for nitrate-nitrogen.   The graphs for Platessa oblongella, however, both have a few outliers.   I have seen P. oblongella in a few situations where I did not expect it – I remember finding it in the Halebourne, a stream draining heathland around Aldershot and Bagshot in Surrey, where the water was well buffered (mean alkalinity: 61.3 mg L-1 CaCO3) and nutrient concentration were high (mean total oxidised nitrogen: 4.01 mg L-1; dissolved phosphorus: 0.25 mg L-1) and Carlos Wetzel and colleagues note some other anomalous records from the literature in their paper (cited in my earlier post), including a few from high conductivity and even brackish environments.   So we should treat these plots as indicative of the ecological preferences rather than definitive.

Distribution of Odontidium mesodon and Platessa oblongella (including P. saxonica) to nitrate-N and dissolved phosphorus in UK streams.   Vertical lines indicate threshold values that should support high (blue), good (green), moderate (orange) and poor (red) ecological status classes.  See Annex 2 for more explanation.

The final pair of plots show how the relative abundance of these two species changes over the course of the year.  These plots show the months when each taxon is abundant, by the standards of that taxon.  Because Platessa oblongella tends to be very numerous in samples, the threshold for this taxon (the 90th percentile of all records) is higher than that for O. mesodon.   This reveals a very clear pattern of O. mesodon thriving in Spring whilst P. oblongella is abundant throughout the year, but with a slight preference for summer and autumn.  We need to reconcile these patterns with the observations in A tale of two diatoms that show that P. oblongella is associated with thinner biofilms than O. mesodon and try to work out whether season is driving the patterns or whether the seasonal patterns are the manifestation of other forces.   My suspicion is that P. oblongella is a classic pioneer species but also has a low-growing prostrate habit which means that it should be resistant to heavy grazing, which may confer an advantage in the summer and autumn when grazers are most active.  However, I may be getting ahead of myself, as we are in the process of analysing data on grazer-algae interactions in the River Ehen and Croasdale Beck that may throw more light on this.  There are clearly more layers to this story yet to be revealed …

Distribution of Odontidium mesodon (i.) and Platessa oblongella (j., including P. saxonica). The solid lines represent relative sampling effort (i.e. the proportion of samples in the dataset collected in a particular month) and the vertical bars represent samples where the relative abundance of taxon in question exceeded the 90th percentile for that taxon (20% for P. oblongella/P. saxonica and 5% for O. mesodon).

Reference

The dataset used for these analyses is that used in:

Kelly, M.G., Juggins, S., Guthrie, R., Pritchard, S., Jamieson, B.J., Rippey, B, Hirst, H & Yallop, M.L. (2008). Assessment of ecological status in UK rivers using diatoms. Freshwater Biology 53: 403-422.

Annex 1: Odontidium mesodon’s preference for still or standing water

As I included a graph showing the preference of Platessa oblongella / P. saxonica for still or standing water in “A tale of two diatoms …”, I have included a similar graph for Odontidium mesodon here.   I have not included any data from the streams that flow into Ennerdale Water’s north-west corner in this graph as this would give a distorted picture.  To date, I have only seen a single valve of O. mesodon during analyses of 14 samples from these streams but I have not yet sampled these in spring which, as the graph above shows, is the time when O. mesodon is most abundant.   Like Platessa oblongella, O. mesodon is predominately a species of running, rather than standing waters.

Differences in percentage of Odontidium mesodon in epilithic samples from Ennerdale Water and associated streams.  Data collected between 2012 and 2018.

Annex 2: notes on species-environment plots

These are based on interrogation of a database of 6500 river samples collected as part of DARES project.  Vertical lines show UK environmental standards for conditions necessary to support good ecological status: blue = high status; green = good status, orange = moderate status and red = poor status.  Note that there are no environmental standards for alkalinity and the vertical lines show a rough split of the gradient into low alkalinity (“soft water”: < 10 mg L-1 CaCO3), low/moderate alkalinity (³ 10, < 75 mg L-1 CaCO3), moderate/high alkalinity (³ 75, < 150 mg L-1 CaCO3) and high alkalinity (“hard water”: ³ 150 mg L-1 CaCO3).

pH thresholds are for clear water (see UK TAG’s Acidification Environmental Standards.  The corresponding thresholds for humic waters are lower (high/good: 5.1; good/moderate: 4.55; moderate/poor: 4.22; poor/bad: 4.03).

Phosphorus thresholds are based on UK TAG’s A Revised Approach to Setting WFD Phosphorus Standards.   Current UK phosphorus standards are site specific, using altitude and alkalinity as predictors.  This means that a range of thresholds applies, depending upon the geological preferences of the species in question.  The plots here show the position of boundaries based on the average alkalinity and altitude measurements in the DARES database.

Note, too, that phosphorus analyses use the Environment Agency’s standard measure, which is unfiltered molybdate reactive phosphorus.  This approximates to “soluble reactive phosphorus” or “phosphorus as orthophosphate” in most circumstances but the reagents will react with phosphorus attached to particles that would have been removed by membrane filtration.

Nitrate-nitrogen: There are, currently, no UK standards for nitrates in rivers.  Values plotted here are derived in the same way as those for phosphorus (see “This is not a nitrate standard”)

 

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.

References

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.

Algae behaving selfishly …

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

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

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

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

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

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

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

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

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

References

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

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

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

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

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

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

 

Change is the only constant …

The diatoms I saw in my sample from the littoral of Lake Popovo (described in the previous post) reminded me of an assemblage that I had seen in another lake which, apart from its location, has much in common with Popovo. This lake is Wastwater, in the western part of the English Lake District (see “The Power of Rock …”).  Like Popovo, it is situated in a remote a region of hard volcanic rocks and, as such, has very soft water and is subject to few of the pressures to which most of our freshwaters are subject.  The photograph above shows me sampling Wastwater in about 2006 (more about this photograph, by the way, in “A cautionary tale …”).

I wrote about Wastwater when I was writing my book Of Microscopes and Monsters, the precursor of this blog.   I focussed, in particular, on an experiment that my friend Lydia King had performed as part of the research towards her PhD.  Her previous work had established that there were relationships between the types of algae that she found in lakes in the Lake District and the amount of nutrients that they contained.  She also saw that the types of algae she found depended upon how acid or alkaline the water was.  But the water chemistry only explained a part of the variation in the algae and now she wanted to find out about the variation that was not explained by this.   In particular, she wanted to know how much of the variation was due to the way that the algae interacted with each other.

Lydia’s experiment involved putting clay pots into the shallows at the edge of Wastwater and then watched how the algal communities changed over the course of six weeks.  She also examined small parts of the pots at extremely high magnifications using a scanning electron microscope.   These micrographs, and subsequent conversations with her, had inspired some of my early paintings and I returned to this subject several times, finally producing a series of three pictures that showed changes in the algae over time.

The microbial world of the littoral zone of Wastwater after two weeks of colonisation showing unidentified small unicellular blue-green alga,  unidentified small unicellular green alga; thin filaments of Phormidium,  Achnanthidium minutissimum and Gomphonema parvulum.

The first of these shows the surface of the plant pot after being submerged in Wastwater for two weeks.   You could think of this as a patch of waste ground that was, at the start of the experiment, bare of vegetation.   If we watched this patch over a number of weeks, we would notice some plants appearing: scattered stalks of grass, perhaps some rosebay willow herb, dock or plantains. A gardener might dismiss these as “weeds”, although this term has no ecological meaning but ecologists prefer to think of these as “pioneers”: plants adapted to colonising new habitats, growing quickly (which might mean producing lots of seeds in a short space of time or producing rhizomes or runners) and covering the ground.  This same process has taken place on Lydia’s plant pot in Wastwater: the “weeds” in this case are scattered thin filaments of the blue-green alga Phormidium, the diatoms Achnanthidium minutissimum and Gomphonema parvulum plus a number of spherical green and blue-green cells that she couldn’t identify.   Such is the scale that we are working at that this open landscape still contains about 92000 cells per square centimetre.

The microbial world of the littoral zone of Wastwater after three weeks of colonisation.   The composition is similar to that in the previous figure but the density of cells is greater.

When she came back a week later, much of the empty space had been infilled; there were now about 300,000 cells per square centimetre.  These mostly belonged to the same species that she had found the week before.  The difference is that they are now rubbing up against each other and this has some important consequences.  All plants need light and nutrients to grow and algae are no exceptions.   Sunlight provides the energy for photosynthesis but now, at week three, the density of algae is such that there is a chance that some of the light will be intercepted by a neighbouring cell.   The total amount of sunlight that filters through the water to the pot surface is already much lower than that available at the lake surface; now it has to be shared out between many more cells.   At this point, properties such as fast growth rates that helped our pioneers to colonise the plant pot become less relevant, and it is algae that are better adapted to capturing the limited light that will survive.

So when Lydia came back to Wastwater after six weeks, she saw a very different community of algae on her pots.   There was still a lot of Achnanthidium minutissimum, but rising above these was the elegant art deco shape of Gomphonema acuminatum (also found in Lake Popovo) which, importantly for our story, grows on a long stalk.  There are also cells of “Cymbella affinis” (the correct name at the time that Lydia was working but see comments in the previous post about the nomenclatural history of this species).   This, too, grows on a long-stalk, the better to grow above the Achnanthidium and other pioneers.   If we continue to use the analogy of a patch of wasteland, then it has now reached the point where it has been invaded by shrubs such as hawthorn and blackthorn.   However, in a terrestrial habitat this would happen two or three years after the first pioneers had arrived, not six weeks as Lydia had observed for the algae.   She also found the diatom called Tabellaria flocculosa which forms filaments.  These often start out loosely-attached to the substratum but more often break free and become entangled around the other algae.   In our “wasteland” analogy, these would be the brambles.

The microbial world of the littoral zone of Wastwater after five weeks of colonisation.  Gomphonema acuminatum, “Cymbella affinis” and Tabellaria flocculosa have now joined the assemblage seen in the two earlier dioramas.

The experiment finished shortly after this, terminated when the apparatus was overturned.  Whether by a wave or by vandalism, Lydia will never know but this event is, itself, a metaphor for the harsh world in which benthic algae have to survive.  In real life, the many cobbles in the littoral zone will be rolled by wave action or, as we have seen in other posts, invertebrate grazers could have removed much of the “shrubbery”, leaving a “pasture” composed of the tough, fast-growing species such as Achnanthidium minutissimum to dominate samples.   The “successions” we see in the microscopic world not only take place much more quickly than those in the macro world, but they also rarely have a stable “climax”: just a brief pause before the next onslaught from the physical, chemical and biological processes that shape their existence.

References

King, L., Barker, P. & Jones, R.I. (2000). Epilithic algal communities and their relationship to environmental variables in lakes of the English Lake District. Freshwater Biology 45: 425-442.

King, L., Jones, R.I. & Barker, P. (2002). Seasonal variation in the epilithic algal communities from four lakes of different trophic state. Archiv für Hydrobiologie 154: 177-198.

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.