Comparing algae on a summer’s day …

I wrote about the effect of the long period of low flow in the River Wear a few weeks ago (see “Summertime Blues …”) and have, now, completed two dioramas depicting the state of the river in the main channel and in a filamentous algae-dominated backwater.  The first of these is dominated by free-living green algae, either single cells or coenobia (see note at end), which is a big contrast to the situation I recorded two months earlier when the assemblage was dominated by diatoms, with patches of filamentous green algae (see “Spring comes slowly up this way” and “A question of scale”).

I sent a small sample of the Wolsingham biofilm to Dave John for his opinion on the green algae, and he sent back a list with twenty one different green algae that he had found.  Fortunately, this confirmed my own original list, with Keratococcus bicaudatus, Scenedesmus, Desmodesmus and Monoraphidium all featuring.   He also commented that Keratococcus is hard to differentiate from Chlorolobium (which is also in his list) and that most of the green alga on his list are usually considered to be planktonic (Keratococcus and Chlorolobium are exceptions) although, as my earlier post suggested, these definitely formed a distinct biofilm on the surface of stones this year in the River Wear.

A diorama showing the biofilm in the River Wear at Woslingham, July 2018.   You can see coenobia of Demodesmuss communis (centre), Scenedesmus sp. (left) and Coelastrum microporum (right – half tucked behind a mineral particle, along with single cells of Keratococcus bicaudatus (upright cells) and Monoraphidium.  There are also some cells of Achnanthidium minutissimum on short stalks in the foreground and a cluster of Fragilaria gracilis cells in the background.

There seems to be little hard evidence on the habit of Keratococcus and Chlorobium apart from references to a preference for benthic habitats.   I have drawn them as upright cells, drawing on their similarity in form to Characium, for which there is better evidence of an upright habit (although Characium tends to grow on other algae, rather than on hard surfaces).  Whereas I often have a strong sense of the three dimensional arrangement of organisms within benthic biofilms, so little has been written about the preferences of these green algae that, apart from the Keratococcus, I have had to show them as a jumble of cells and coenobia across the picture frame.

The second diorama depicts the tangle of filamentous green algae that I found in the pools beside the main channel.  As I mentioned in my earlier post, these are species that I do not normally find at this site and are here, I presume, due to the long period of unusually warm weather and low flows.   One difference between these communities and that captured in my first diorama is that there is a more obvious organisation of the constituents here: the Cladophora filaments, though appearing as a tangle to us, form the foundation upon which epiphytes grow directly, but also around which Melosira filaments are entangled, rather like the lianas in a tropical rain forest.   The quantity of diatoms around the Cladophora is so great that their brown pigments completely mask the Cladophora’s green cells but note how the density of Cocconeis cells reduces towards the tips – the youngest parts of the filaments.

Depiction of filamentous algae growing in the margins of the River Wear at Wolsingham in July 2018, showing epiphytic Cocconeis pediculus and entangled Melosira varians.

There have been some recurring themes in my posts this summer: one is that UK rivers have been behaving quite differently to previous years, due to a combination of low flows (more accurately, a lack of the scour associated with high flows) and warm, well-lit conditions.   The low flows have also resulted, to some extent, in rivers becoming more physically heterogeneous, with side-pools and silty areas developing distinct assemblages of algae quite different to those encountered in the main channel.   Sometimes, the sum of these effects is for rivers to look less healthy than is usually the case.

The Wear at Wolsingham is one of those sites that I like to think I know well, having visited the location so many times over the past 30 years.  It is reassuring, in a rather humbling way, to know that it still has the capacity to surprise me.

Dave’s list of green algae from the Wolsingham biofilm, July 2018

Desmids
Closterium moniliferum
Closterium acerosum
Cosmarium botrytis
Cosmarium venustum
Staurastrum striatum

‘Chlorococcalean’ algae
Acutodesmus dimorphus
Coelastrum astroideum (very small and atypical)
Coelastrum microporum (very small and atypical)
Chlorolobion braunii
Desmococcus olivaceum (subaerial species)
Desmodesmus communis
Desmodesmus subspicatus
Keratococcus bicaudatus
Monoraphidium arcuatum
Monoraphidium contortum
Monoraphidium griffithsia
Monoraphidium irregulare
Scenedesmus arcuatus
Pseudopediastrum boryanum
Tetradesmus obliquus
Tetraedron minimum

Note

A coenobium is a colony in which the cell number is fixed at the time of formation and not augmented subsequently.   Coenobia are particularly common in the Chlorococcalees.

Transitory phenomena …

Fieldwork in the River Ehen has been an unusually pleasurable experience over the past few months, even to the extent of abandoning waders altogether and wearing just a thin pair of neoprene beach shoes and shorts as I worked.   Curiously, there were few obvious signs of the prolonged period of low flow here, but that is partly due to the pumps installed by United Utilities to keep the river running whilst the lake was drawn down (see “Life in the deep zone …”).   I did, however, find some intriguing green patches on fine sediments at the margins.

Most of the bed in this part of the river consists of much coarser sediments than these which are, I suspect, silt and sand deposited on the occasions when Ben Gill (which joins the Ehen immediately below Ennerdale Water) is flowing.   Current velocity is lower at the edges of the river, allowing fine sediments to settle out and create temporary sandbanks.   One decent spate will be all that is needed, I suspect, to wash much of this downstream.  However, there has not been a period of prolonged high flow for several months and there is, as a result, a thin green mat of algae growing on the upper surface of this sediment.

Mats of Oscillatoria on fine sediments beside the River Ehen just downstream from Ennerdale Water, August 2018.   The total length of the mats in the left hand photograph is about one metre. 

I scraped up a small sample to examine under my microscope.  I was expecting to see the broad filaments of the cyanobacterium Phormidium autumnale which I often find at a site about five kilometres downstream (see “’Signal’ or ‘noise’?”) but what I saw was much narrower filaments, some of which were slowly gliding forwards and backwards.   These belong to a species of Oscillatoria, a relative of Phormidium that is common in the plankton.  A few species, however, do live on surfaces and can, as I could see in the Ehen, form mats.  I have, in fact, described a different mat-forming species of Oscillatoria (O. limosa) from the River Wear close to my home (see “More from the River Wear”) and this, too, had been favoured by a long period of warm weather and low flow.   The filaments in the River Ehen were much narrower – just a couple of micrometres wide – and had relatively long cells (two or three times longer than wide) but, in other respects, they clearly belonged to the same genus.

Microscopic views of Oscillatoria filaments from the River Ehen, August 2018.   The upper photograph was taken at medium magnification (400x) and the lower image was taken at 1000x.  The constant motion of the filaments means that it is not possible to use stacking software to obtain a crisp image.  Scale bar: 10 micrometres (= 1/100^th of a millimetre). 

The motion that I could see is thought to be due to a layer of tiny fibres (“microfibrils”) which wind around the inner layer of the cell wall in tight spirals.   Movement is caused by waves that are propagated along these fibres, meaning that the filament actually rotates as it moves (though this is almost impossible to see with a light microscope).   The filaments can move either towards or away from light, depending on the intensity, at a speed of up to 11 micrometres per second (that’s about a millimetre a day or, for any petrolheads who are reading, 0.00004 kilometres per hour).  This allows the filaments can adjust their position so that they are neither in the dark nor exposed to so much light that they are likely to do damage to their photosynthetic apparatus (see “Good vibrations under the Suffolk sun” for more about this).   The result is that filaments will tend to converge, Goldilocks-style, at the point where light conditions are “just right”.  You can see some sediment particles settling on the top of the mat in one of the images and we can expect the filaments to gradually adjust their positions, incorporating these particles, over time.

Last year, I wrote about Microcoleus, a relative of Oscillatoria, which formed mats on saltmarshes and explained how this could be the first stage of colonisation of damp habitats by plants (see “How to make an ecosystem”).   We are seeing the same processes happening here, but the life expectancy of these mats is much lower.  They may well be gone next time I visit, depending on how the Cumbrian climate behaves over the next couple of weeks.   They are transitory phenomena, here today and gone tomorrow but, like the subjects of some of my other recent posts, particularly favoured by the long period of settled weather that we have enjoyed over recent weeks.

Reference

Halfen, L.F. & Castenholz, R.W. (1971).  Gliding motility in the blue-green alga Oscillatoria princeps.  Journal of Phycology 7: 133-145.

Note: you can read more about how the heatwave has affected fresh water in the Lake District in Ellie’s MacKay’s recent post on Freshwaterblog

Two-faced diatoms …

Back in March I reflected on the challenges involved in discriminating species of Gomphonema (see “Baffling biodiversity …”).   That there were several species in the sample which prompted the article was indisputable; that some of those species were, individually, quite variable was also clear.  The former issue I resolved, to some extent, by reference back to Hutchinson’s “Paradox of the Plankton” but the latter was harder to explain.

Part of the problem stems, I suspect, from the reliance on morphology to characterise species.  We assume that, because a group of organisms share a set of visible characteristics, then they must also share genes which determine those characteristics and that, in turn, implies a common ancestry.   Turning that assumption on its head, we assume that groups of microscopic algae that appear different to each other belong to different species.   However, a dog lover might point out that Chihuahuas and Great Danes look very different but are, in fact, the same species.   One of the challenges of those of us who study algae is deciding just how much variation in form is typical within a species, and at what point differences are such that they represent more than one species.

Gomphonema sarcophagus from Pitsford Water, Northamptonshire, showing Janus cells.  Photographs by Ingrid Jüttner.  Scale bar: 10 micrometres (= 1/100^th of a millimetre).

So what should we make of the diatom valves in the image above?   The valve outlines and breadths are similar but the striae densities are so different that we might think that they belong to two separate species.   However, I recently stumbled, by chance, on a 1998 paper by Stacy McBride and Robert Edgar which discussed the topic of “Janus cells”.  Janus, you may remember, is the Roman god of time and is depicted with two faces, one looking back to the past and the looking to the future. His name has been appropriated, in this context, to describe diatoms that have frustules comprising two valves with different characteristics.   A few genera show consistent differences between the two valves – in Cocconeis and Planothidium, for example, one valve has a raphe whilst the other does not – and there are also differences in striae densities between the raphe and rapheless valves.   The term “Janus cell” is applied to diatoms where there are marked differences between the two valves but this is not a fundamental characteristic of the species or genus.   So, in the example above, we see some forms with much denser striae (11-13 in 10 mm) than others (7-8 in 10 mm).

We don’t know, from just looking at variability in populations, that this is not polymorphism within the species, in much the same way that some humans have attached ear lobes and others do not.   But, as diatom populations grow in number by repeated divisions of single cells, we can assume that most are clones of a small number of genotypes and, therefore, that the differences are due to ontogenetic variation.   What is interesting here is that this variation seems to create two distinct outcomes – coarsely or finely striated valves.  Some have suggested that such variation may be determined by differences in environmental conditions; however, the co-existence in a single population argues against this.

Gomphonema, as I have mentioned in earlier posts, is a genus that challenges taxonomists.  And, because ecologists depend upon taxonomists to give them a means of sorting diatom valves and frustules into meaningful categories, the environmental signals we get from Gomphonema species are often quite confused too.   The possibility of encountering Janus cells just throws one more curve ball into the mix.

Reference

McBride, S.A. & Edgar, B.K. (1998).   Janus cells unveiled: frustular morphometric variability in Gomphonema angustatum.   Diatom Research 13:293-310.

Round, F.E., Crawford, R.M. & Mann, D.G. (1990).  The Diatoms: Biology and Morphology of the Genera.   Cambridge University Press, Cambridge.

Life in the colonies …

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

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

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

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

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

Reference

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

Getting started with microscopy

I talked about algae last week at an event organised by Durham Wildlife Trust (part of the publicity for The Natural History of Upper Teesdale) and I promised them a post about how to get started in microscopy and, more specifically, to start discovering more about algae.  I have illustrated the post with some of Chris Carter’s spectacular images of algae to whet your appetites for exploring the world of freshwater algae …

Broadly speaking, the natural historian wants a microscope for one of two tasks: to make small things bigger or to make invisible things visible.   There is not really a sharp dividing line between these two categories, as the illustrations of Cladophora filaments in “Summertime blues …” show.   You might start out looking at a handful of green slime pulled from your garden pond, but then you might see smaller algae growing on those filaments that you want to examine too.   The good news is that you should be able to get hold of a reasonable microscope with the capacity to magnify up to 400 times for about the same outlay as a digital SLR camera.   That should let you see all but the smallest algae.   If you are sure that your interests lie mainly in “making small things bigger” then you should consider a low power dissecting microscope (these are probably the best way of introducing children to microscopy, as there is a smooth transition between the tangible but small object that has piqued their interest and the larger, more detailed image that they see when they peer through the microscope’s eyepieces).

However, even though a basic microscope need not cost a fortune, good microscopes are expensive so my advice to a beginner is to search out a rerfurbished second hand microscope.  In north east England, I can recommend JB Microscopes but Google should help you find dealers elsewhere in the country.   A reasonably local supplier is necessary because you should really try out a microscope before you buy.   There are reputable mail order suppliers (e.g. Brunel Microscopes) but I would not want to spend a large sum on a piece of equipment that I had not had a chance to use first.

A colony of the diatom Meridion circulare.  The image at the top of the post shows desmids from the genus Micrasterias.   Both photographs by Chris Carter. 

If you are on a limited budget, I suggest you go for a good basic microscope with the option to fit a camera at a later stage.  It is possible to take a reasonable photograph by pointing a digital camera (or even a smartphone) down the microscope’s eyepiece and it is better to put up with the shortcomings of these images than to sacrifice the quality of the microscope itself.

Once you have your microscope, you will also need slides, coverslips, forceps, some plastic Pasteur pipettes, a couple of needles, a scalpel and some collecting tubes.  You can buy all of these from Brunel Microscopes and NHBS, both of whom cater for both the amateur and professional markets.   They also sell boxes of prepared slides, which are a good way to get some experience at using a microscope.

The microscopic world generally lacks the type of user-friendly well-illustrated identification guides that help us identify wild flowers, birds, butterflies and so on.   Most books are aimed at the academic market and are, consequently, expensive.   If you want to get started with freshwater algae, one useful resource is this guide to the larger algae found in rivers: RAPPER_manual_version1.7_May2016.  It was produced to accompany a method for rapid assessment of streams and rivers and, as the journey towards formal publication has stalled, I am happy to make it available here.

Hydrodictyon reticulatum, the water-net, photographed by Chris Carter.  500 mm (micrometres) is half a millimetre.

Useful websites include AlgaeVision and the Diatom Flora of Britain and Ireland.  As most freshwater algal genera are found throughout the world, Diatoms of North America is also a useful resource.

The Freshwater Biological Association have affordable booklets on the identification of desmids and diatoms and there is an AIDGAP key, too, for freshwater diatoms.   The latter is badly in need of updating but, people assure me, is still useful for beginners.

There are plenty of other online resources, but l would recommend visiting the website of the Quekett Microscopical Club, a long-established group of enthusiasts whose interests span the whole realm of natural history and optics.   www.microscopy-uk.org.uk is also worth a visit.   Both websites will help you as you start your explorations of the hidden worlds of nature.