Lecture 18 Animal Diversity 6

Protostomes to deuterostomes

The last and very diverse group of arthropods mentioned in last lecture are the crustaceans –
“aquatic” mandibulates [33.39]. They all have shells at least partially thickened with calcium
carbonate – some are sessile and relatively simple (e.g. barnacles), others are mobile with a
dazzling array of specialised appendages (e.g. lobster; cleaner shrimps [see Life Episode 8
Creatures of the Deep on BB resources, additional movies]). Not all live in water, e.g.
pillbugs/woodlice, but all require at least moist conditions. All are in the ecdysozoa – they have to
grow new exoskeleton before moulting [see spider crabs in Life Episode 8 and on display in a case
visible from the road in our museum]. (Anyone keen on crustaceans will enjoy the TED talk on
Mantis shrimp http://www.ted.com/talks/sheila_patek_clocks_the_fastest_animals.html)

The aim of this lecture is to finish off the Protostomes (and therefore to explain the one remaining
major distinction in the animal group, that between Protostomes & Deuterostomes), to review some
of the diversity of the echinoderms and to introduce chordates – animals with a “backbone”.

The last group in the protostomes is the tardigrades – (tardus – slow; gradus – step). The
tardigrades are sometimes called water bears or moss piglets and are very successful, >1,000 spp
named (see http://www.youtube.com/watch?v=XOxXHJGKsYI for the movie from the lecture). These simple
animals are mostly terrestrial and live in the water film around mosses and lichens. They have 4
pairs of short unsegmented legs which bear 4-8 simple claws with which they cling to vegetation or
sand and creep about (slow steps) One reason for considering this group is that these animals
have parallels with the first plants on land – they too are famous for their ability to enter state of
suspended animation (cf. some mosses). If dried gradually, their water content decreases from
85% to 3%, movement stops and they can survive for a century or more in temperature extremes,
ionising radiation and oxygen deficiency. (see previous website and https://www.youtube.com/watch?
v=SUC0_HjNFBs). Add water and they rehydrate and start moving in minutes. Tardigrades have also
been sent into space by man, and shown to have remarkable powers of both survival in extreme
conditions and recovery afterwards, see http://tardigradesinspace.blogspot.com/ and some people
consider the survival of tardigrades in space to support an extra-terrestrial origin of life. For more
information (and to prepare for the next eLearning module) see
http://www.lab-times.org/labtimes/issues/lt2009/lt02/lt_2009_02_10_11.pdf and the article on Water Bears
on the BSR area of resources.

Deuterostomes differ from the protostomes (e.g. molluscs, annelids and arthropods) in 3 important
ways (handout; 32.9). (1) the way their embryos divide (radial vs spiral) and determinate (vs
indeterminate – each cell’s fate is not determined as soon as it is formed). (2) the way the body
cavity (coleom) forms (no need to know details) (3) the fate of the blastopore – the mouth forms
from this in protostomes (protos = first; stoma – mouth); in deuterostomes (deuteros = second) the
anus forms from this and the mouth is formed from a secondary opening – the one that forms near
the opposite end of the animal. Both deuterostomes and protosomes are bilaterally symmetrical
animals (Bilateria), but it is not at first obvious when considering the adults of most of the familiar
simplest forms of Deuterostomes alive – echinoderms, which include sea urchins and sea stars
[handout]. [33.40]

Echinodermata – (echinoderm – means “spiny skin”). >7,000 species, a very successful group. A
calcareous endoskeleton is found in all, e.g. as part of proper spiny skeleton (as in sea urchins and
sea stars). Adults of many of this phylum appear radially symmetrical, but the larvae are clearly
bilaterally symmetrical. They are amongst the most abundant marine animals, especially at extreme
depth. Members of the sea cucumbers don’t look much like the rest of the echinoderms but some
are quite spiny. They feed by extending mucus-covered tentacles then putting them into mouth to
collect particulate matter. Some can evert their digestive tract, and some can also evert their
respiratory organs when threatened [see handout] – respiratory organs can form sticky strands and
some contain toxins – can be (slowly!) regenerated. They are good at healing wounds and it has
been discovered that they use a similar mechanism to regenerate parts – so biomedical
researchers are very interested in finding out the molecular mechanism behind this [Biomed Sci
students might like to see http://www.sciencedaily.com/releases/2007/10/071017194617.htm].
Their relatedness to sea urchins and sea stars becomes clear from their locomotion. This is via
tube feet, which are in rows along the body, and are part of a water vascular system. This is easiest
to understand from sea stars [handout, 33.40 and movie [similar close up of tube feet can be seen
in “Monster worm and sea star frenzy” in resources; additional movies on BB ]) which have an
interconnected network of canals. The system is open to the outside via a perforated plate
(madreporite} that leads to the water outside the animal. A ring canal around the centre connects
with radial canals, branching from each of which are hundreds of hollow muscular tube feet filled
with fluid and which is continuous with the rest of the system. Each tube foot consists of bulb-like
portion, which can inflate the foot portion and allow it to extend and contact the substratum. Each
foot portion has a sucker that can grip, and has muscles in its wall that can contract and force fluid
back into the bulb portion. This allows (slow) locomotion, ‘righting’ of the animals if upside down
(e.g. earlier movie on Killer Alga Caulerpa, where urchins were tested on time taken to “right”) and
can also help during feeding – the tube feet can detect prey and then help the sea star prize open
bivalves such as clams, as the suction exerted by the tube feet can reach 30g – most starfish can
evert their stomachs, to envelop their prey (or into the opening of a clam) and digests the contents
[movie from Life Episode 8. Creatures of the Deep on BB additional movies; same process in
“Monster worm..” too]

The mouth of sea urchins is underneath the animal – equipped with an Aristotle’s lantern – a
muscular device powering 5 sharp teeth [Hickman 23.20, handout, movie and see the good
animation in http://www.asnailsodyssey.com/LEARNABOUT/URCHIN/urchFeed.php#s] that graze
on algae and other organic material – eg holdfasts of Macrocystis. Urchins can cause huge
environmental problems if predators (eg sea otters) not present or are removed [see Planet Earth
clip “Sea Urchin Barrens” http://www.bbc.co.uk/programmes/p0038tcc].

Sexual reproduction is also very simple – eggs and sperm are shed into water and fertilisation is
external. Very few echinoderms show parental care but as the gametes easy to collect and handle
we know more about development, biochemistry and molecular biology of embryos of sea urchins
than almost any other system . [Developmental Biology students should see Chapter 47 in
Campbell] The larvae are clearly bilaterally symmetrical – reorganisation is required to generate a
radially symmetrical juvenile.

Chordata. These are the most familiar animals – those with a rod-like, semi-rigid body of cells
enclosed by fibrous sheath that extends the length of body. Chorda – cord, and all members of this
phylum share a notochord (noton – back). [34.3; handout]. There are both invertebrate and
vertebrate chordates, all are linked by four trademarks of the phylum Chordata.

1. Notochord
2. Dorsal tubular nerve cord
3. Pharyngeal pouches
4. Post-anal tail

Notochord is longitudinal flexible rod located between digestive tube and nerve cord. This provides
support either for muscles or skeleton. In most vertebrates this becomes replaced by vertebrae.

Nerve Cord develops from plate of ectoderm that rolls into tube dorsal to the notochord. In humans,
this forms our brain and spinal cord.
Pharyngeal pouches. The region behind the mouth is called a pharynx. In aquatic chordates the
pouches are flow-through. This allows water entering the mouth to exit again without going into the
rest of the digestive tract. This can generate suspension feeding devices but most are modified for
gas exchange.

Post-anal tail. In most non-chordates the digestive tract extends nearly the whole length of the
body. The tail is a structure “added” to the body beyond the end of the digestive tract, its evolution
allowed more efficient propulsion in water (esp. for fish). The tail is now evident in humans only as
a series of small vertebrae at the end of our spinal column (coccyx), but most other mammals have
waggable tail as adults.

We’ll get onto familiar vertebrates at the end of chordates, but first, more primitive examples.
Urochordates - sea squirts [movie]. Urochordates means “tail chordates”, but the adults don’t have
tails (or anything else much really!) they are sessile u-shaped suspension feeders [34.5 and movie].
Their chordate nature is only clear in the larvae (which are constructed just like the “model
chordate” in the handout diagram – 34.5).

One other animal is very like the idealised chordate in Fig 34.3 - the Lancelet ( Branchiostoma is the
name usually preferred to Amphioxus, which is the one used in the tree of life, but no need to
remember either!) [34.4 and handout], a small animal, lives in sand at sea bottom in coastal
regions. These animals can reach huge density (>5000 m -2 in Tampa Bay, Florida). These animals
have tentacles at their anterior end and are filter feeders (trap particles in mucus on cilia in
pharyngeal slits) – no jaws, no brain case (cranium) and no real brain – just a vesicle at the end of
their notochord, which is where segmented muscles are anchored, seen easily in these animals as
they are translucent. Such animals may well resemble the ancestral form that gave rise to the rest
of the chordates (movie from Life on Earth).

Craniata - all the members of the group do have a braincase (cranium), but some lack vertebrae.
Hagfish have a rudimentary cranium and endoskeleton of cartilage (not labelled in the diagram on
the handout; movie) and are crucially important as bottom-dwelling scavengers [34.9]. They are
environmentally important as agents of recycling of nutrients from dead/dying animals. They are
often found at huge depth – they have a keen sense of smell/touch, and can detect carcasses on
the sea bed from tens of kilometres away. The slime glands can generate several litres of slime in
<1 min – this repels or even suffocates other scavengers or predators – slime is employed by many
animals (handout; and perhaps see the following, which does have some good science in, but I am
not sure I would recommend the hagfish slime scone recipe!
http://twistedphysics.typepad.com/cocktail_party_physics/2010/08/killer-snot-monsters.html ).
Hagfish feed by rasping away chunks of flesh using keratinised plates on their tongues
in a pincer-like action [handout] (keratin = really useful structural protein – we have
it in hair and nails). They are able to tie knots in their tails and pass it forward
along their body for extra leverage [handout and movie] because they have no stiffened
bony skeleton or vertebrae.
BIOL10511Lecture Key learning outcomes Key words
18 Understand the key Protostome
differences between Deuterostome
protostomes and Chordate
deuterostomes, and the Craniate
key characteristics of
chordates.

Question for (PASS) discussion

Can you remember two animals that inject toxins into their prey before they feed on them? How
comes those animals are not affected by the toxins in their food?
What are the advantages of a supporting rod along the back of an animal, and what are the
disadvantages of stiffening that?