Lecture 16 Animal Diversity 4

Molluscs to arthropods - organs and appendages.

Case study of Caenorhabditis elegans.

Triploblastic animals – the aim of this lecture is to review how the third layer of tissue in such
animals led to diversification of organs and increasing sophistication of form; to review the
diversity of reproductive strategies in metazoans; to review modifications that allowed
metazoans to colonise terrestrial habitats.

As we saw last time, molluscs all have the same body plan – based on a muscular “feeding
foot”. We did not mention reproduction last time, but many molluscs are hermaphrodites, and
mating in such animals allows both partners to be fertilised and thus maximises the
reproductive output (one reason why slugs and snails are such an agricultural and horticultural
problem). Many have internal fertilisation – the sperm is delivered into the female reproductive
tract, and thus parental care.

The most complex, and the most intelligent molluscs are the cephalopods – “head-foot”
animals. The foot is modified and concentrated in the anterior region, and part of it takes the
form of a funnel (siphon) for expelling water from the mantle that surrounds the animal e.g.
Nautilus, cuttlefish, squid, octopus. The only living cephalopod with an exterior protective shell
is Nautilus, supremely well adapted to their mode of life as highly mobile predators, see
http://www.youtube.com/watch?v=QMFqV4SJLWg (essential to understand how they move and
hunt, and if you are interested to find out how they optimise their position in the water, and are
able to be so active, or interested in evolutionary history of animals, see also “Nautilus:
Chambers of secrets” New Scientist 05 April 2008 http://www.newscientist.com/article/mg19826501.700-
nautilus-chambers-of-secrets.html).

Cephalopods have the largest eyes of any animal (up to 25cm) - they are “camera eyes” like
our own and allow these animals to locate predators (handout) as well as to locate prey and
mates very effectively, and they exhibit some remarkable behaviours – e.g. see Life episode
mentioned in last lecture (Creatures of the Deep, on BB) and New Scientist “The secret
language of cuttlefish” 26 April 2008 (including an excellent movie entitled “do you speak
cuttlefish”) http://www.newscientist.com/article/mg19826531.000-the-secret-language-of-cuttlefish.html)
Many cephalopods live at extreme depth but come to the surface to mate [movie] in a
synchonised spawning. The simplest have external fertilisation – e.g some squid (movie
showed males hanging on to females in order to fertilise the eggs externally as they are laid,
whereupon the parents disappear back into the depths – no parental care). Molluscs include
both monoecious (single sex) and dioecious (hermaphrodite) animals (e.g. snails are
hermaphrodite). Mating can be a long and even elegant affair – see the leopard slugs in “Life
in the Undergrowth” - http://www.bbc.co.uk/sn/tvradio/programmes/lifeintheundergrowth/video.shtml . In the
most advanced molluscs, fertilisation is internal, eg sperm are delivered into female giant
squid during copulation (to see a preserved giant squid, visit the Natural History Museum in
London, you need to book a spirit collection tour; see http://www.nhm.ac.uk/kids-only/life/life-sea/giant-
squid/). Some of these highly intelligent animals also exhibit elaborate parental care behaviours,
e.g. “Broody squid carry their eggs in their arms” New Sci 14 December 2005
http://www.newscientist.com/channel/earth/deep-sea/dn8462-broody-squid-carry-their-eggs-in-their-
arms.html and, the extreme situation in the BBC “Life” Episode 8 programme (octopus parental
care) on BB.

Cephalopods are protected almost as comprehensively as mammals by UK government

guidelines on animal experimentation. Their intelligence and complex behaviour reflects the
fact that cephalisation has been taken further than in the gastropods – they have a
recognisable brain (concentration of ganglia) e.g. the second section of this movie:
http://www.youtube.com/watch?v=V6mtZSzYeM0&feature=related; see also “Defensive tool use in a
coconut-carrying octopus” Current Biology, Volume 19, Issue 23, R1069-R1070, 15 December 2009; (see the
movie in supplemental material - to go straight to full text version
http://www.cell.com/current-biology/fulltext/S0960-9822(09)01914-9 )

More information on intelligence, of likely interest to neuroscientists, can be found in: “Octopus
Behavior, Intelligence, Learning, Personality, Problem Solving & Consciousness” SLA
Biomedical and Life Sciences Blog August 04, 2010
http://sla-divisions.typepad.com/dbio/2010/08/octopus-behavior-intelligence-learning-personality-
problem-solving-consciousness.html (and coconut-use movie therein). Lastly, some octopus are
exceptionally good at mimicry, see: http://www.youtube.com/watch?v=H8oQBYw6xxc
Annelids – ‘little rings’ a.k.a. bristle worms. These are segmented animals with a body cavity
(coelom) with specialised organs, lined from one end all the way to the other in mesodermal
tissue. This “tube within a tube” organisation allows organs to develop with the luxury of
suspension. The non-compressible fluid of the cavity functions as a hydrostatic skeleton. The
cavity provides space for organs, including sex organs (usually of both types) so mating usually
involves exchange of sperm.

Earthworms are of huge environmental importance – see BSR article in resources (250 tonnes
of casts produced per hectare per year – aerates soil; carries debris and dead leaves
underground where they are eaten, or broken down by decomposers; carries seeds and spores
both down from and up to surface). Segments are partitioned off from each other by septa, this
allows just one part of body to move at a time (handout; 33.24). In non-segmented animals any
fluid changes or muscular contractions are transmitted all the way through the body, whereas
segments allow different compartments to behave differently (movie). This also allows for failure
– if one segment is damaged, the problem is not transmitted to the neighbouring segments so
function may be maintained by them. Earthworms can also reproduce asexually by breaking
into separate sections and regenerating the missing parts.

Annelids are also sometimes known as ‘bristle worms’ as most bear bristles (setae) that anchor
their bodies during locomotion. Many are active burrowers and setae also help prevent the
worm being pulled or washed out of their burrow. Not all annelids have internal fertilisation like
the earthworm. Christmas tree worms release their gametes into the water (ie no parental care)
but also show synchronised spawning (thus enhancing the chances of male meeting female
gametes). These annelids make protective tubes, and all that divers/snorkellers see will be the
anterior, which is the “Christmas tree” that functions as gas exchange (cf. gills), feeding
apparatus, and as a reproductive conveyor belt.

Nematodes (roundworms) (>25,000 species named). Most widespread of all animals. They
exhibit ecdysis (moulting of the exoskeleton – an outermost supportive/protective layer) – the
exoskeleton of roundworms is not rigid but tough and flexible – it is mostly the structural protein
collagen.

Case study nematode: the soil-dwelling Caenorhabditis elegans (see the following for a movie
of the development of an adult worm from an egg http://www.ifg.tu-bs.de/Schnabel/Celegans01.mov).
The origin and lineage of all 959 cells in its transparent body have been traced from zygote to
adult, the ‘wiring diagram’ of its nervous system (all neurons and connections) has been
mapped, the programmed cell death that results in the final animal has been documented in
detail and its genome was one of the first to be sequenced. It is very easy to culture in the lab,
using simple media, and it breeds quickly. A vast suite of mutants have been generated and
some have cast light on the process of a huge range of diseases e.g. muscular dystrophy,
others have been used in studies of ageing, and see handout for just a few of the latest uses for
this model organism. Interestingly, although it has been in lab cultivation for decades, only
relatively recently has its role in the natural world been investigated (Environmental Scientists
and Zoologists might like to see “”The natural history of Caenorhabditis elegans” Current Biology
20, 2010 http://www.cell.com/current-biology/fulltext/S0960-9822(10)01168-1

Arthropoda
(“Jointed feet”) easily the most successful animal group...3 out of every four animals is an
arthropod. The success is usually attributed to their segmented body - each segment can
become specialised for a different function – as can the jointed appendages – modifications
include walking, feeding, sensory reception, copulation, defence (see handout; 33.30 for the
“blueprint” of an arthropod). They have a versatile exoskeleton: layer of protein enriched with
chitin - strong and relatively impermeable to water (allowed colonisation of land). Extensive
cephalisation (most sensory organs concentrated at anterior). Efficient respiratory system –
internal surfaces specialised for gas exchange in terrestrial arthropods.

Chelicerates – e.g. scorpions, spiders, ticks: most anterior appendages of some members of the
simplest arthropods are pincers (claws) or poison fangs (eg spiders) none has jaws so they
have to crush or liquefy their food.

Spiders Are all predators – they ambush or trap their prey, inject venom using fangs at the ends
of anterior appendages (chelicerae).
 They liquify prey tissue with digestive fluid and then suck up the resultant broth.
 Rearmost appendages include spinnerets – that lay trip wires or make webs to catch
prey using silk (movie)
 Other important factors include book lungs – just like gills stacked in plates contained in
an internal chamber (handout; 33.33) The extensive surface area enhances exchange
of O2 and CO2
 They have internal fertilisation eg male spider spins web, deposits a drop of sperm on it,
picks it up and stores it in special cavities of his palps. When mating he inserts a palp
into the female and dispenses the sperm (same movie as above).
 Mating sometimes involves males and females very different in size from each other
 often preceded by elaborate courtship displays

Ticks are mostly blood sucking parasites – with highly sophisticated barbed mouthparts. Once it
has found a suitable host, a tick attaches itself by means of sharp skin cutters and a barbed headpiece
(hypostome) – handout. The hypostome also serves as the food channel that directs blood into the tick’s
mouth – see “The secret life of ticks” in BSRs in lecture resources (written by Dmitri Logunov in our
museum; who has also written a nice article on jumping spiders you can find on BB). In order to feed
successfully a tick has not only to attach itself tightly, but also to stay undetected by the host. Tick saliva
plays a crucial role. It suppresses the local immune and inflammatory responses – anti-inflammatories are
just one of the many biomedical applications of ticks (handout).

Ticks carry some serious diseases in their saliva, so the BSR article on BB is highly
recommended reading for anyone going out and about in the countryside, or on field trips in
dense vegetation!

BIOL10511Lecture Key learning outcomes Key words

16 Understand how Cephalopod
 molluscs and Hermaphrodite
arthropods reproduce. Cephalisation
Understand advances Arthropod
that led to colonisation Exoskeleton
of land. Chelicera

Questions for (PASS) discussion.

Do you consider the ability to mimic other organisms a sign of intelligence?

Why might it be a rational response for humans to be scared of spiders?