BIOL10511 lecture 4

Prokaryotes to eukaryotes
(Fairly simple, mostly small things, to more complex small and bigger things)
Case study of euglenoids and dinoflagellates.

The emphasis in the last lecture was the metabolic and structural versatility of the
prokaryotes. The aim of this lecture is to finish off the prokaryotes and consider
how they are thought to have contributed to the emergence of eukaryotes, and
also to show how they and some of the smallest eukaryotes are also of great
environmental and biomedical importance.

Cyanobacteria fix more than ½ nitrogen fixed by micro-organisms. “Algae” as a

whole fix over ½ the carbon fixed on earth. I mentioned earlier that some
cyanobacteria can move, others are also able to make sure they position
themselves in perfect place to photosynthesise – those that can are seen to have
small bright spots within their cells under a light microscope. From transmission
electron microscopy we know that these things comprise many small cylinders
which contain an atmosphere less dense than the surrounding cytoplasm and are
resistant to pressure – gas vesicles [handout]. When the sun is shining it causes
a high rate of photosynthesis in the cells in the top layers of a body of water (e.g.
lake). High rates of photosynthesis mean lots of sugar molecules are produced
and this causes an increase in cell turgor pressure. When the turgor pressure
gets sufficiently high, it can cause gas vesicles to collapse. That causes the
organisms containing them to sink. This puts them in a position where less light
penetrates, so there is a low rate of photosynthesis. The low rate of
photosynthesis causes a decrease in cell turgor pressure. This means that when
new vesicles form they don’t collapse and the cells can rise. These inclusions in
the cells allow cyanobacteria that have them to optimise where they are with
relation to the light. This may not be right at the surface of the water, it might be
several centimetres below that. Each organism can optimise where it is in the
water table, it’s not at the mercy of wind currents, water currents and convection
currents within the water body. Only some cyanobacteria contain gas vesicles,
those that do have a distinct advantage over those that don’t.

As introduced in last lecture, during prolific growth cyanobacterial “blooms” can

form – this can be an excellent food source (e.g. Spirulina – last lecture), but see
also New Sci 9th Nov 2002, p8 “Bug suspect in bird deaths”
http://www.newscientist.com/article/mg17623680.400-bug-suspect-in-bird-deaths.html and
more recently in news: http://www.bbc.co.uk/news/uk-england-cambridgeshire-
23518616

So cyanobacteria can generate severe environmental problems, even the non-

toxic ones, as when they are at high density other plants are robbed of light and
can start to rot, which starts to affect the animal life in the water. So quite quickly
you can get very large bodies of water being very smelly, hostile and even toxic
[see BSR Vol 15 issue 3 33-36,2003, “Blue green algal blooms” – see BB].
EUKARYOTES
First the “Protists” (according to the outdated 5 Kingdom system).

For about 2000 million years before organisms we might call plants, fungi and
animals were distinguishable, our planet was inhabited solely by the prokaryotes
we have covered so far and a group of organisms informally known as protists.
[see Chapter 28 for a LOT more information than is required for this unit]. The
protists are the most diverse of any group of Eukaryotes – the aim of the rest of
this lecture is to give a general overview and then to focus on just 2 groups to
illustrate the difficulty of distinguishing groups of organisms at this level (and the
wisdom of the 3 domain system we are using).

The handout [similar to figure 25.9 in the eighth edition of the textbook] gives a
plausible pathway for the formation of cells with mitochondria and plastids, via
engulfment of hetero- or autotrophic prokaryotes. Just as prokaryotes pass
genes to each other today (via conjugation etc – see lecture 2) engulfed
prokaryotes (endosymbionts; endo – inside, sym – together, bios – life) have
exchanged some genes. Advocates of genetically manipulated (GM) organisms
stress that the correct way to picture the relationships between groups or
organisms in the 3 domains of life is as shown on the handout. By transferring
genes between organisms we are merely continuing something which has been
going on since life began (and see the New Scientist article linked in the
“Additional interesting stuff” forum on the Discussion Board).

There is good evidence that engulfment of organisms occurred very much more
often than once. To take just one example, Euglena [28.7], the chloroplasts of
which have more than the usual 2 layers of membrane around their chloroplasts.
The interpretation of how this arose is shown on the handout [28.2] – plastids
derived from endosymbionts that were eukaryotic algae that in turn contained
plastids derived from cyanobacteria. There are many other variations on this,
accounting for some of the bewildering taxonomy of the photosynthetic
eukaryotes (there is no need to learn about groups other than those featured in
lecture notes).

Along with ultrastructural diversity comes nutritional and structural diversity – and
euglenoids have many “ways to make a living”. In the light, chloroplasts allow
Euglena (“good eyeball” – so named because of the conspicuous eyespot) to be
autotrophic [28.7]. A single flagellum pulls the cell through the water towards the
light using a lasso-like motion [handout]. If cells are in darkness, Euglena can
live as a heterotroph by absorbing organic compounds through its permeable
proteinaceous covering (called a pellicle) – not a rigid cellulose cell wall as in
(other) plants, but a flexible covering that also allows Euglena and relatives to
move through relatively solid substrata eg mud. [movies – see e.g.
http://www.youtube.com/watch?v=J3sgoy22Ij4 for a good movie of Euglena
movement using the flexible pellicle and some nice darkfield images, but you
need to go back to the Oscillatoria movie for a good view of Euglena using
flagellar movement – around 17 seconds in http://www.youtube.com/watch?
v=lCm_UZd8IKk].

Euglenoids without chloroplasts can ingest particles of food. So the euglenoids

include autotrophs (that can act as heterotrophs – as in some cyanobacteria
mentioned last lecture) and heterotrophs. They appear in textbooks both as
protozoa and as algae - these 2 terms are often used, but clearly have no
taxonomic significance. This morphological and metabolic plasticity means that
sometimes Euglena can form problematic booms.

The handout [28.2] shows the endosymbiotic events assumed to have given rise
to the main lines of photosynthetic eukaryotes. Green and red algae from
primary endosymbiosis; euglenoids and dinoflagellates, secondary.
Dinoflagellates are an example of a large group of important protists called the
Chromalveolata [handout] – meaning coloured alveolates (the “brown tide”
organism mentioned in the article on the handout is another organism in this
group).

Dino probably comes from the Greek word dinos meaning whirl (not deinos –
terrible), and “flagellate” means having flagella. Most are covered in plates of
cellulose armour [handout], which protects them and makes them rigid, relatively
robust and able to withstand the shearing forces as they move (very rapidly!)
through the oceans. In flagellate dinoflagellates, a girdle flagellum runs around
the middle of the cell, which causes the cells to spin [28.9]. A posterior flagellum
drives them forwards. So these organisms whirl through the water in a helical
pathway [You can find a movie of a dinoflagellate swimming in the CD that
comes with the textbook if you bought it and there is a short movie at
http://is.gd/mwSlia]. They have chromosomes which are always condensed
[handout]. This condition is referred to as mesokaryotic – nothing like prokaryotes
and not much like other eukaryotes, with some really unique cell biology.

Dinoflagellates are very environmentally important, mainly as the smallest

primary producers in aquatic environments. But when parts of the oceans of the
world turn red, they are described as “red tides” [handout] – this is caused by a
bloom of dinoflagellates. These are considered such a problem that the media
even warns of impending red tides (e.g. see
http://www.treehugger.com/files/2010/02/significant-red-tide-bloom-forcast-for-new-england.php (some
other short articles that might be of interest about algae are cited at the end), and
again, more locally: http://www.bbc.co.uk/news/uk-scotland-highlands-islands-
19332572 As the last link indicates, some red tides and blooms are dangerous
because of the neurotoxins they contain. Organisms with simpler neural systems
than our own, eg shellfish, fish, eat them with few or no ill effects. “Paralytic
shellfish poisoning” (after eating “dodgy shellfish”) can be caused by
dinoflagellates. Fish contaminated with neurotoxins from dinoflagellates can also
kill humans, or at least make them extremely unwell.
The toxins are used by the dinoflagellates as predators, as they are also
mixotrophs – capable of both autotrophy and heterotrophy (like some
cyanobacteria and euglenoids, so they are also sometimes described as
animals). [If you are feeling brave see if you can understand a primary research
paper: “A dinoflagellate exploits toxins to immobilize prey prior to ingestion” January 19,
2010, doi: 10.1073/pnas.0912254107
http://www.pnas.org/content/early/2010/01/14/0912254107.full.pdf+html but this is
strictly very optional reading, for anyone who wants to find out what the research
papers are like that the text book and lecturers base their information on].
Dinoflagellate toxins are listed in Schedule 1 of the Chemical Weapons
convention.

Many dinoflagellates are also bioluminescent. Why they produce light when
disturbed was not understood until recently [handout and see
http://www.youtube.com/watch?v=bCNjXaMPZxw]. (more next lecture on this).

Many anemones, and hydroids, such as corals, contain these organisms. It has
been estimated that there are over thirty thousand dinoflagellates in a cubic
millimetre of coral.

BIOL10511Lecture Key learning outcomes Key words

4 Understand how gas vesicles Gas vesicles
work. Bloom
Understand how prokaryotes are Endosymbiosis
thought to have given rise to Flagellum
eukaryotes via endosymbiosis. Euglenoid
Understand the environmental and Dinoflagellate
biomedical importance of Bioluminescence
euglenoids and dinoflagellates. Red tide

Questions for (PASS) discussion

Is an organism that can switch between photosynthesis (autotrophy) and

heterotrophy a plant or an animal?
Why should lecturers refer to primary research papers (and is the stuff in them
likely to be examined)?