The annual Algal Training Course in Durham always has a field trip out to Cassop Pond, a small pond at the foot of the Permian Limestone escarpment in County Durham that has featured in a few of my posts over the years (see “A return to Cassop”). This year, the group came back with some samples from the pond’s margins bearing a suspension of green dots just visible to the naked eye which, when examined under the microscope, turned out to be the colonial green alga Volvox aureus. These are spherical, with the cells at the periphery, joined together by thin strands of protoplasm. The smaller colonies were scooting about, propelled by the pairs of flagellae borne by each of the cells that constitute the colony, whilst the larger ones (mostly “pregnant” with one or more daughter colonies) were sessile.
Volvox aureus colonies just visible to the naked eye in a drop of water from Cassop Pond, July 2019. The drop is 13 millimetres across.
Colonies of Volvox aureus (each bearing daughter colonies) from Cassop Pond, July 2019. Scale bar: 50 micrometres (= 1/20thof a millimetre).
A close-up of part of a colony of Volvox aureus from Cassop Pond, July 2019. Scale bar: 20 micrometres (= 1/50thof a millimetre).
Watching a Volvox colony swimming around under the microscope is a beguiling experience, but its movement is not random. Consider: there may be a 1000 or more cells in the larger colonies, each with two flagellae. If all beat their flagellae at random, the colony would not get anywhere, as the force in one direction would be cancelled out by forces in all other directions. But Volvox colonies do actually move with intent. Look closely at the individual cells in the photos below and you will see that each has a red-coloured eye spot (the light-detecting organelle actually lies beneath the red layer, which acts as a filter). People with more patience than me have noticed that the eye spots in different parts of the colony differ in size, suggesting a level of organisation that may not be immediately apparent. We also know that the daughter colonies tend to form at the posterior end of the colony (assuming “posterior” and “anterior” in a spherical colony are defined by the direction of travel) and also that only a small number of cells (larger than the others) are responsible for the division that produces these.
In theory, a spherical object is going to offer less resistance and so sink faster than an object of the same size that had a greater surface area : volume ratio. This should mean that they are not able to stay in the light-rich surface layers where they can photosynthesise and grow. In practice, Volvox colonies are able to adjust their position by using their flagella but this requires them to pump some of the energy they have obtained from photosynthesis into the flagella’s motors. Another advantage in Volvox’s favour is a relatively low density of the colony as a whole. The individual cells are separated by strands of protoplasm which creates a lattice through which water can penetrate, so the overall density of the colony is closer to that of the surrounding water than would be the case if the cells were tightly packed.
Volvox is most often found in the summer in relatively nutrient rich lakes, where nutrients are sufficiently plentiful to support a rich crop of algae. A motile colony that is not too dense is well-placed to adjust its position to stay in the surface layers and harvest the sunlight. Moreover, the size of the colony probably means that it is too big for the filter-feeding zooplankton that grazes on the algae. At the same time, however, Volvox begins to experience some of the problems associated with multicellular life (see references in “The pros and cons of cell walls …”). As large multicellular organisms ourselves, a nuanced discussion about the pros and cons of multicellularity may seem to only have one possible outcome. However, Volvox inhabits a world where plenty of single-celled organisms thrive and where a colonial lifestyle offers a small competitive advantage. It means that it is quite happy drifting around at the time of year when many of us would like nothing better than to don swimming trunks and soak up some sun in a local pool. Study algae for too long and you end up realising that only losers need to evolve.
Cells from a Volvox aureus colony from Cassop Pond, July 2019. You can see the red eye-spots in some of the cells in the left-hand image (bright-field) whilst the protoplasmic strands joining cells together can be seen in the right-hand image (phase contrast). Scale bar: 10 micrometres (= 1/100thof a millimetre).
Canter-Lund, H. & Lund, J.W.G. (1995). Freshwater Algae: Their Microscopic World Explored. Biopress, Bristol.
Reynolds, C.S. (1984). The Ecology of Freshwater Phytoplankton. Cambridge University Press, Cambridge.