Cassop Pond in May

A month on from the phycological debauchery that I wrote about in Promising Young Algae the Spirogyra flocs that covered quite a lot of the surface of Cassop Pond have disappeared.  With sexual reproduction over, the zygotes, I presume, have sunk to the bottom of the lake, where they will lay dormant until next Spring.   I searched around the margins of the pond, but only found a few wisps of Spirogyra hanging around some cattle hoofprints in the shallow water on the eastern side of the pond.   This, however, proved to be a different type of Spirogyra altogether, with broader and squatter cells than the main constituents of April’s flocs.  

The green tinge in this hoofprint is filaments of Spirogyra.   The picture at the top of the post shows Cassop Pond in May 2021: note the absence of flocs compared with the situation in April. 

Swimming around amongst the Spirogyra filaments were a number of very active green cells of Euglena.   Euglena is a genus that has not featured much in this blog over the years (see: “A visit to Loughrigg Fell”) but it is a genus closely associated with Cassop Vale, with 13 of the 36 species recorded from Britain and Ireland recorded from this location.  The story behind this richness is that there were no active experts in the Euglenophyta at the time when the Freshwater Algal Flora of the British Isles was being compiled.   Instead, a Polish expert, Konrad Wołowski, was invited to contribute and, to help him do this, he was taken on a tour of locations that Dave John and Brian Whitton, the editors, thought would be likely habitats.  Cassop Pond is well known to Brian and is conveniently located near Durham, so it was an obvious location.  As a result, it is probably the hot spot of recorded diversity for this genus in the UK but that is more due to the idiosyncrasies of biological recording than to anything about this location over many others that particularly favours Euglena

That said, cattle hoof prints are known to be a good location for Euglena and relatives, and I wrote about a relative of Euglena that I found in a puddle in Teesdale (see: “Puzzling puddles on the Pennine Way …”.  A hoofprint or a puddle is, from an alga’s point of view, a temporary pond and so long as you have a plan in place for when this dries up, it represents a potential habitat.   Cattle, as we have already seen, are allowed to graze on the reserve so there are plenty of damp hoofprints within which Euglena and relatives can thrive.  That’s also true of many other nature reserves around the country.  The one missing ingredient at all of those is naturalists within an inclination to search them out …

Spirogyra filaments from Cassop Pond, May 2021.  Scale bar: 20 micrometres (= 1/50th of a millimetre).

Reference

Wołowki, K. (2010).  Euglenophyta.  pp. 181-239.   In: Freshwater Algal Flora of the British Isles (edited by John, D.M., Whitton, B.A. & Brook, A.J.). Cambridge University Press, Cambridge.

Euglena sp. from Cassop Pond, May 2021.  Scale bar: 10 micrometres (= 1/100th of a millimetre).  

Some other highlights from this week:

Wrote this whilst listening to:   The soundtrack to The Pursuit of Love, available via BBC Sounds.   

Cultural highlights:   Emily Mortimer’s adaptation of Nancy Mitford’s The Pursuit of Love, available on the BBC iPlayer

Currently reading:  Pat Barker’s Noonday, the final part of the Life Class trilogy.

Culinary highlight: homemade digestive buiscuits.

Promising young algae …

Spring has arrived in Cassop Vale.  Leaves are appearing on many of the trees and the ground vegetation has the green flush of a new beginning.   More importantly, the herd of emo-fringed highland cows have been moved away, to give the plants more chance of flowering, and there is some warmth in the sun in the middle of the day.

From my point of view, the biggest change since I was last here is the appearance of an extensive floc of green algae covering much of the pond’s surface.   I had a hunch, from their appearance, that these would be predominately Spirogyra, but was not expecting the sight that greeted me when I put a small piece of a floc under the microscope. 

Flocs, predominately Spirogyra, in the margins of Cassop Pond, April 2021.

I find Spriogyra and its relatives quite regularly on my travels, but usually in the vegetative state.  It is relatively unusual to find them as they undergo sexual reproduction (see “Fifty shades of green …”).  But there was plenty of evidence of this process (termed “conjugation” in Cassop Pond’s green flocs.  There were plenty of vegetative filaments, each about 20 micrometres wide and with a single helical chloroplast.  But there were also many ellipsoidal zygotes apparent.   When I looked more closely, these were inside filaments which were linked to an adjacent filament by a narrow tube.   What started out as an early morning natural history trip has turned out to be the algal equivalent of Saturday night on Newcastle Quayside.   

For those of you unused to dating, Spirogyra style, here is a quick guide.   First, put on your best helical chloroplast (two or more, if you are daring), then head out to find a partner amongst the many other filaments in your particular floc.   Little is known about Spirogyra’s preferences, but we can assume that many species are not heterosexual, so don’t be shy: sidle up to any filament you fancy.   He/she/it might well play hard to get at first, so maybe you need to drop a hint.  Make sure your potential date gets a whiff of your aftershave (that’s what I assume “hormonal interactions between the paired filaments” means).  If he/she/it gets the hint, then you can indulge in a little mutual meiosis to get yourselves into the mood.    

Spirogyra from flocs in Cassop Pond, April 2021.   a. vegetative filament; b. two filaments undergoing sexual reproduction with zygotes in the lower filament.   Narrow filaments of Aphanizomenon gracile are also present.   Scale bar: 20 micrometres (= 1/50th of a millimetre).  

Now we’ve got that all-important emotional (okay … hormonal) connection, it is time to get physical.   An embarrassing bulge appears on the side of your filament but, fortunately, a similar one should appear on the side of your date’s filament at about the same time.   Eventually, these fuse to form a tube that links you both together.  The correct term for this is the “copulation canal” which is as frank as it is alliterative (it could also be called a “tupping tube”, I guess?). The protoplast of both cells now contracts and one (the “boy”, for want of a better analogy) crawls, amoeba-like, through the tube and fuses with the “girl” protoplast to form a zygote.  That’s as far as our frisky filaments in Cassop Pond have got.  If our phycological peep-show continued for longer, we would see the green zygotes gradually become brown in colour as thick, resistant walls grew around them, and the cell contents were processed into starch and lipid-rich food reserves.   They would then sink to the bottom of the pond and rest, dormant, until conditions were ripe for its germination.

Features of Spirogyra conjugation: a. a vegetative cell in one of the two aligned filaments; b. conjugation canals developing between the aligned filaments; c. a zygote.  Scale bar: 20 micrometres (= 1/50th of a millimetre).  

Why here, why now?   Nitrogen limitation has been quoted as one of the triggers for conjugation and the presence of a nitrogen-fixing cyanobacterium (Aphanizomenon gracile) plus nitrogen-fixing diatoms (Epithemia– see “Working their passage”) in the pond at the same time lends support to this hypothesis.  Also, the yellow-green appearance of the flocs is also a hint that they may be nitrogen-limited.   However, there are also reports of conjugation happening on a predictable annual pattern in some locations.  The two possibilities are not mutually exclusive, we should remember.  

Meanwhile, on dry land, there are plenty of other plants getting down to the complicated business of reproduction too.   We saw goat willow (Salix caprea) and hazel (Corylus aveana) as well as lesser celandine (Ficaria verna) in flower, and leaves of primroses yet to bloom.   You can read more about those here.   Just remember, when enjoying the sight of spring flowers, that the botanical bacchanalia takes place in less obvious ways in the water too.

Some other highlights from this week:

Wrote this whilst listening to:  Horses and Easter by the Patti Smith Group (see below).   And a 1977 BBC “Sight and Sound in Concert” recording of Jethro Tull, which I remembered seeing when it was first broadcast.

Cultural highlights:   The film Black Bear – a rather dark and challenging, but ultimately rewarding, film.

Currently reading:  Just Kids, by Patti Smith.  Best read with Horses and Easter as a soundtrack.  The geographer in me also reads it with a map of New York to hand, as it is a book with a very strong sense of place.

Culinary highlight:.our local Indian restaurant makes a rather good lamb shank, cooked in aromatic spices which, with basmati rice and a side order of bhindi, is just about unbeatable.

Natural lenses …

The photograph above is as about as far from Andreas Gursky’s careful constructions, described in the previous post, as it is possible to get.  It is a close-up of a green algal floc Heather noticed whilst on a walk around a local nature reserve.   I guess it fits the general description of “decisive moment” except that it takes a special sort of observer to find any interest at all in such an unprepossessing habitat.

Under the microscope, the floc turned out to be composed of filaments of Spirogyra, with a single helical chloroplast.  Members of this genus (and related genera such as Mougeotia) produce copious mucilage so are always slimy to the touch.  However, this mucilage makes it difficult for the waste gases produced by photosynthesis to diffuse away, leading to the production of bubbles within the mucilage mass.   The interest, today, however, was that these air bubbles are acting as tiny lenses through which it is possible to make out the individual filaments of Spirogyra.

The green floc beside a footpath in Crowtrees local nature reserve from which the other images in this post were derived. 

I should add the caveat here that the photograph was taken with the “super macro” facility of our Olympus TG2 camera but the end-product is, nonetheless, impressive.   It also offers us an insight into the world of the very earliest microscopists.  Anton van Leuwenhoek’s microscopes consisted of a metal plate which held a tiny sphere of glass which acted as a convex-convex lens capable of up to 266x magnification to a resolution of little  more than a micron (1/1000th of a millimetre) (follow this link for more details).  To give an idea of what he might have seen with this, the right hand image below used 400x magnification.

That, however, only tells us part of the story of Anton van Leuwenhoek’s genius.   Whilst we should not underestimate the skill required to make the lenses and their mounts, the other essential element is curiosity.   Curiosity is, itself, multifaceted: in a few weeks we will probably make a trip out to an old quarry where we know we will find several species of orchids, and maybe some excursions to locations new to us but where others have reported interesting assemblages.  That’s one type of curiosity.  However, simply looking harder at the habitats all around us involves a different type of curiosity: a recognition that there is more to know even about things we think we already know about.   The former broadens our experiences, the latter deepens them …

The algal floc at Crowtrees local nature reserve in close-up: left: an extreme macro view of a single bubble from the image at the top of the post and, right: filaments of Spirogyra photographed under the microscope.  Scale bar: 20 micrometres (= 1/50th of a millimetre).

Concentrating on carbon …

On the other side of Ennerdale Water I could see plenty more submerged stones, all covered with green filaments but these belonged to different genera to those that I wrote about in my previous post.   Both are genera that we have met previously – Mougeotia, which has flat, plate-like chloroplasts which rotate around a central axis in order to control its rate of photosynthesis – and Spirogyra.  When light levels are low, Mougeotia’s flat chloroplast is perpendicular to the light in order to capture as much energy as possible, but in bright light it rotates so that the plate is parallel to the direction of the light, in order to slow the photosynthesis mechanism down and prevent internal damage (see “Good vibrations under the Suffolk sun” for another approach to this problem).

However, too much sunlight is the least of an alga’s problems in the Lake District.   This post looks at a different challenge facing freshwater algae and our starting point is the spherical nodules, “pyrenoids”, that you should be able to see on the chloroplasts of both Mougeotia and Spirogyra in the images below.   Photosynthesis involves a reaction between water and carbon dioxide to make simple sugars (turning fizzy mineral water into “pop”, in other words).   A submerged alga does not have a problem obtaining the water it needs, but what about carbon dioxide?   Gases are not very soluble in water, so this presents a much bigger problem to the algae.   Explaining why also presents a big problem to a blogger who conscientiously avoided physics and chemistry from age 16 onwards.  Here goes …

Mougeotia from the littoral zone of Ennerdale Water, April 2017.  Scale bar: 20 micrometres (= 50th of a millimetre).

The concentration of a gas in a liquid depends upon the concentration of that gas in the surrounding atmosphere.   As far as we know (and this is still an area of contention amongst geologists), concentrations of carbon dioxide in the deep past were much higher than they are today, in part because there were no land plants to suck it out of the atmosphere for their own photosynthesis.  So the earliest photosynthetic bacteria and, subsequently, algae, lived in water that also had higher concentrations of carbon dioxide.   As land plants spread, so the carbon dioxide concentration in the atmosphere dropped as they used it to fuel their own growth.  As a result, carbon dioxide concentrations in the water also dropped, thus depriving the algae of an essential raw material for photosynthesis.

However, carbon dioxide is not the only source of carbon available to aquatic organisms.   There is also carbon in many rocks, limestone in particular, and this can mineralise to carbonate and bicarbonate ions dissolved in the water.  Aquatic plants can get hold of this alternative carbon supply via an enzyme called carbonic anhydrase.   By concentrating the carbonic anhydrase activity in a small area of the chloroplast, the algal cell can boost the activity of the Rubisco enzyme (which evolved to function at a higher concentration of carbon dioxide).   This whole process is one of a number of forms of “carbon concentrating mechanism” that plants use to turbocharge their photosynthetic engines (see “CAM, CAM, CAM …” on my wife’s blog for more about a terrestrial version of this).

A two-chloroplast form of Spirogyra from the littoral zone of Ennerdale Water, April 2017.  Scale bar: 20 micrometres (= 50th of a millimetre).

Pyrenoids are widespread amongst algae, though a few groups (notably red algae and most chrysophytes) lack them.   Cyanobacteria (blue-green algae) use an organelle called a “carboxysome” for a similar purpose.   The only group of land plants with pyrenoids are the hornworts, relatives of mosses and liverworts.   About half of all hornworts have pyrenoids and a recent study has suggested that the ability to form pyrenoids has evolved up to five times in this group during their evolution.   The appearance of pyrenoids in distinct evolutionary lineages of algae also suggests that there may have been several evolutionary events that precipitated their formation.  And, it is important to stress, some algae which lack pyrenoids have alternative methods of concentrating carbon to enhance Rubisco activity.

So let us end where we started: in the littoral zone of Ennerdale Water on an April morning, gazing at a fine “fur” of filamentous algae clinging to the submerged rocks.   Back in October last year, I talked about how Ennerdale fitted into a pattern of increasing productivity of Cumbrian lakes first noticed by Pearsall in the early part of the 20th century (see “The power of rock …”).   Now we can start to understand that pattern in terms of basic biochemical processes: getting enough carbon from a combination of atmospheric carbon dioxide and the surrounding rocks for Rubisco and the other photosynthetic enzymes to convert to sugars.   In Ennerdale Water, one of the least productive of the Cumbrian lakes, we can see these algae during the winter and spring because the amount of biomass that those biochemical reactions produces is still just ahead of the amount that grazing invertebrates such as midge larvae can remove.  In a month or so, the grazers will have caught up and the rock surfaces will be, to the naked eye at least, bare.

Rubisco is the enzyme whose gene, rbcL, we use for molecular barcoding, subject of many recent posts (see “When a picture is worth a thousand base pairs …”).  My early desire to avoid physics and chemistry at school translated into as little biochemistry as possible whilst an undergraduate and, over the past few-years, I’ve developed a frantic urge to catch-up on all that I missed.   Just wish that those lectures explaining the Calvin cycle had been a little less … tedious …

References

Giordano, M., Beardall, J. & Raven, J.A. (2005).  CO2 concentrating mechanisms in algae: mechanisms, environmental modulation, and evolution.   Annual Review of Plant Biology 56: 99-131.

Villareal, J.C. & Renner, S.S. (2012).  Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years.  Proceedings of the National Academy  of Science of the USA 109: 18873-18878.