BIOL10511 Lecture 20 Partners and robbers. 1.

The aim of the next two lectures in the course is to bring together the preceding lectures,
which reviewed biodiversity largely by considering organisms in isolation, so they will also
contain quite a bit of revision.

Symbiosis (Sym - together bios - life) merely means “living together” so includes
parasites and organisms both loosely and tightly in association. The aim of this and the
next lecture is to revise some of the examples we have already considered, to introduce
some important partnerships and to consider how they contribute to biodiversity. We will
start with an emphasis on mutually beneficial partnerships – mutualistic symbioses.

Mutualistic symbiosis is an extremely widespread phenomenon, and involves every type

of organism with every other. In almost every environment there are clear advantages to
both partners – such as the luminescent bacteria that are protected within the ventral
surfaces of squid that allow the squid to pretend to be moonlight so that they can descend
through the water onto their unsuspecting prey at night [handout], and there are many
other examples of animals using the light from luminescent bacteria in some of the movies
I have uploaded on “additional movies”, the angler fish is another example [ see
http://www.youtube.com/watch?v=fMTHrvskSW8 zoologists might also like to see the
really cheesy explanation of angler fish reproduction that follow the fishing lure
explanation on http://www.youtube.com/watch?v=Lsmxs0uDXMo] and see 27.19). (All are
examples of animal-bacterial symbiosis)

Animal-animal symbioses include the relationship

between anemones (Cnidarians with powerful
stings) and anemonefish (clownfish), where the
anemone provides a predator-free environment
for the fish and the fish defend the anemone
against predators (mucus coating on the fish
seems to prevent nematocysts discharging and
harming them) and provide additional oxygen at
night [handout]. [Environmental Scientists and
Zoologists might like to read more about this and a range of other fascinating animal-
animal symbioses in Coral Reefs 24:67-73 2005
http://springerlink.metapress.com/content/9xbchpgr5gj64afl/fulltext.pdf]

Coral reefs are a classic example of how symbiosis allows organisms to survive in
environments neither partner could colonise alone. Especially when nutrients are limiting
(e.g. in the clear warm waters that host coral reefs), intimate symbiosis can be the only
way to survive [movie]. In this mutualistic relationship between coral animals and
photosynthetic partners there are clear benefits from living together – the algae
(dinoflagellates) obtain protection, CO2 and minerals, the animals obtain metabolisable
sugars and oxygen. Other marine examples seen in the movie include clams (molluscs -
handout), jellies (radiate animals, cnidaria), sea anemones (ditto) and dinoflagellates –
algae. We have covered all these in earlier lectures, and also sea slugs – nudibranchs,
molluscs without shells which feed on chlorophyte algae such as Codium [handout] - the
chloroplasts of which can continue to photosynthesise for many weeks after the alga is
consumed. It has recently been found that some sea slugs not only sequester chloroplasts
but also have algal genes in their genomes, and this allows the animal to live purely by
photosynthesis. See New Sci 24th Nov 2008 Solar powered sea slug harnesses stolen
plant genes http://www.newscientist.com/article/dn16124-solarpowered-sea-slug-harnesses-stolen-plant-
genes-.html,

Many symbioses involve profound changes in the two organisms, e.g. behavioural – giant
clams open wide (exposing their delicate fleshy mantles, something which they would be
unwise to do unless the behaviour was highly beneficial), this allows their
photoautotrophic partners to photosynthesise – which are also altered - they leak up to
80% of their photosynthates; structural - jellies that lack nematocysts (they also have
behavioural modifications – they move so that they are positioned in the sun at the start of
the day, follow the sun across the lake, then move down at night to where mineral
nutrients are more plentiful [movie]); corals formed into “bush” shapes that intercept light
effectively [movie] and dinoflagellates (in anemones) without their cellulose plated armour
or flagella [handout].

There is a huge diversity of terrestrial plant-animal symbioses, e.g. Rattan plant [movie]
and Acacia with ants [handout]. Provision of detachable food bodies (e.g. for larvae)
[movie and handout] and even homes (e.g. hollow thorns) by such plants is rewarded by
behaviour that benefits the plants (e.g. deterrence of herbivores, removal of fungal spores,
destruction of competing seedlings, detachment of tendrils of climbers) [movie].

Another example of mutualism involving ants

is the fungus-gardening ants [see “A bug’s
life”]. The ants remove portions of leaves
from plants, march back to their
underground chambers and “feed” the leaf
portions to their fungi. This can create huge
pathways through tropical forests [handout]
showing the routes followed by millions of
leaf-carrying ants. (An example of an animal-fungus symbiosis).

Another example considered in earlier lectures was Azolla, the water fern [handout].
When the cyanobacterium Anabaena is involved in symbiosis with this fern, almost all of
the cyanobacterial cells become heterocysts. The cyanobacterium becomes heterotrophic
- dependent upon the fern for photosynthate, but the fern benefits from the nitrogen fixed
by the cyanobacterium (so, indirectly does the rice with which the fern is grown (in
nutrient-poor paddies), and there is a direct relationship between amount of Azolla and
rice yield [handout]).

A similar story is found in the cycads, which have established a close relationship with
cyanobacteria in root nodules. Roots that become nodulated are apogeotropic (apo –
away from, geo – earth) – they grow up towards the surface of the soil [handout]. (Both of
these examples are plant-cyanobacterium symbioses).
The two most important terrestrial groups of symbioses in environmental terms are
mycorrhizas and lichens. Here we are dealing with plant-fungus symbioses. We covered
mycorrhizas in earlier lectures (e.g. in relation to earliest land plants - again, first
colonisers of land must have found a low nutrient environment), and present day plants
almost all benefit from enhanced water and mineral absorption of fungi in association with
roots [handout]. Fungi may link entire ecosystems (Plant Scientists, Microbiologists,
Environmental Scientists and anyone interested in ecology might like to read New Sci 26 th
March 2011 46-49 “Heard it on the grapevine: The secret society of plants”
http://www.newscientist.com/article/mg20928051.500-heard-it-on-the-grapevine-the-secret-society-of-plants.html
[Environmental Sciences and Microbiology students might also like to see “The woodwide
web” – Nature 7th August 1997 “The ties that bind” and “Net transfer of carbon between
ectomycorrhizal tree species..”]. Organic carbon is not only channelled into the fungi to
fuel the production of fruiting bodies [movie] but the connections can even benefit
neighbouring plants by providing nutrients as well as a communication network. Some
relationships start as a one-way benefit (e.g. orchid seeds are so tiny that their reserves
are insufficient to fuel germination and growth – seedlings can only survive if they start out
by “robbing” organic carbon from fungi [movie featuring the micropropagation unit in the
Royal Botanic Gardens, Kew]) and only later become photosynthetic and therefore
mutually beneficial.

Lichens are extremely diverse, and especially successful and numerous (over 25,000
species have been identified - lots more than the 13,000 mentioned in the movie or the
17,000 in the text book – see p695) in harsh environments e.g. the Namib desert [movie],
and bare rock faces. Many form keystone “species” (actually 2 organisms – they take
their names from the fungus), such as the “reindeer mosses” that can dominate arctic
tundra – not mosses at all, but they can provide the sole fodder for reindeer during the
winter months. Lichens are often brightly coloured as their algal cells would be damaged
by high light levels [31.23 and see some of the “Field Specimens” outside the Stopford
building].

The main bulk of each lichen is the fungus, which envelops the algal cells in a distinct
layer just under the surface [31.24 and handout] – although the fungal partner may
reproduce via its usual fruiting bodies these only produce fungal spores, which have to
locate their algal partner in order to re-form a lichen. Reproduction of the whole
partnership is achieved by tiny propagules (soredia), which contain both fungal and algal
cells [handout] so when they are blown or broken off, they can grow to form another lichen
in a new location.

BIOL10511Lecture Key learning outcomes Key words

Understand what symbiosis Mutualistic
20 means, and how valuable it
can be in certain Lichen
environments.

Question for (PASS) discussion.

Can the relationship between a sea slug which contains plant genes and that eats algae
and maintains their chloroplasts in a fully functional state be described as a mutualistic
symbiosis?

What stops lichens dominating all terrestrial ecosystems, rather than solely the extremes
of the sub-arctic and the desert?