BIOL 10511 Lecture 14

Animal Diversity 2 – what makes an animal?

Protists to Metazoa
Textbooks tend to imply that the relationship between plants and animals is of over-riding
ecological importance to our planet, but as mentioned in earlier lectures, some highly productive
ecosystems are not based on primary production of plants, but microbes. In the 1970’s,
submersibles allowed the first examination of the deep sea, mid-ocean ridges, 2 miles deep and
28,000 miles long – they represent largest mountain chain on earth [movie]. Vents, where the
earth’s crust comes into contact with the sea, generate water hotter than molten lead (400ºC liquid
water). These hot vents support a rich ecosystem, based entirely on extremophile
microorganisms. Polychaete worms (Pompeii worms [handout]) survive at temperatures that can
exceed 80ºC (Nature 391 545-546 1998 “Worms bask in extreme temperatures”
http://www.nature.com/nature/journal/v391/n6667/full/391545a0.html) mussels (molluscs), crabs
(crustaceans) and tube worms (annelids) (all invertebrates) also contribute to the community
based on bacteria metabolising the sulphides from the vents [handout]. Cold seeps are a similarly
non-plant-based ecosystem, this time the main input for bacteria is usually methane.
Chemoautotrophs provide the primary source of energy for over 500 species of animal; the top of
the food chain is fish (vertebrates) – the animals in these ecosystems include most of the major
clades of Eumetazoan animals we will consider in later lectures [33.2 and handout]. (Eumetazoa -
zoa = animal; meta = later; eu = true). The aim of this lecture is to consider how Eumetazoa
arose from protozoa (proto = first) and reconsider the difficulty of distinguishing ‘animals’ from
plants and fungi.

Just as we think all plants arose from flagellate protist like Chlamydomonas, we think all animals
arose from non-photosynthetic flagellated protist. There are several-celled protozoa such as the
choanoflagellates [32.3] – collared flagellates which are thought to resemble these organisms. As
with the plant kingdom, the assumption is that many cells are better than one – there was
evolutionary pressure for cooperation between cells.

One group of protozoa favoured for the study of multicellularity, which involves communication
and co-ordination between cells, and differentiation of tissues are the cellular slime moulds
(Mycetozoa - “fungus–animals” - they are not fungi, but produce spore-bearing structures that
reminded the folks who first discovered them of fungi) – social amoebae such as Dictyostelium.
These are ecologically important as decomposers – they feed on organic detritus or bacteria using
pseudopodia, like Amoeba. The textbook introduces Dicty [28.25] but my information source for
the following and for the illustrative slides and movies were Chris Thompson (who will expand on
this lecture in his lectures next year) and Professor Cornelis Weijer (see University of Dundee,
Biocentre website).[Genetics and Biochemistry students might like to see his BSR article in lecture
resources for more information]

When food (bacteria) is plentiful, Dicty amoebae live as single cells in the soil. When starved, up
to 500,000 aggregate into a mass. The cells that contribute are usually genetically distinct. This
aggregate can function as a single unit that can form a mound, or alternatively a slug which can
move around, and finally (after about 24h) form a fruiting body with a stalk supporting a mass of
spores that is raised above the substratum. Dispersal and, when conditions are suitable,
germination follow – a new population has formed in better conditions than those experienced by
the previous generation.
This is another system like mating in Chlamydomonas where we have very detailed information
about the events in the process (and can cultivate the organism, which has a host of characterised
mutants, very easily in lab conditions). Aggregation is the result of cyclic AMP production by a
cell. This causes chemoattraction of other cells. The biochemical machinery ensures cyclic
behaviour of each cell, every cell that has been activated acts as chemical oscillator producing
cAMP. It is possible to follow the subsequent fate of individual cells in the mass, which forms a
coherent mound or a ‘slug’, and then the fruiting body using fluorescent proteins. Cells do not
have a fixed position, but move within the ‘slug’ (movie), and although they all start off identical,
they differentiate into two cell types (that become stalk or spores in the fruiting body), the groups
of cells and movement of the slug is also controlled by cAMP. The cells have no fixed fate at this
stage, slugs can regenerate to form two fruiting bodies of normal proportions with the right cell
types when cut in 2 pieces. Injection of cAMP (movie) produces a new tip in intact slugs. [for
most of the short movies from the lecture see http://dev.biologists.org/cgi/content/full/128/22/4535/DC1]
This means that the aggregation of cells can react to predation (e.g. the tip getting eaten) and the
piece left behind can redifferentiate each of the right cell types in the right proportions. The slug
moves around behaves like a multicellular organism but the cells are not connected to each other.

If aggregation happens on the soil surface, the mound goes ahead and makes fruit body. If it is
under the surface, the slug forms and migrates to the surface. The organism senses and reacts to
its environment and can regenerate lost parts. [to see almost the whole life cycle in a movie:
http://www.youtube.com/watch?v=518MhtVBJho]

The life cycle [28.25 and handout] is largely spent in the haploid condition (as in
Chlamydomonas), with only a brief period of the diploid (2n) condition upon fusion of gametes,
with meiosis following immediately. The simplest animals are therefore very similar to the
simplest plants, and Dictyostelium provides clues about how ancestral single celled non-
photosynthetic organisms (protozoa) may have generated the earliest animals - Metazoa.

Sponges, sometimes known as Parazoa (“beside the animals”) are at simplest level of true
multicellular forms. They are not considered “true animals” (Eumetazoa) because the cells are not
organised into tissues, but include amoebocytes (cells like Dictyostelium) and choanocytes (cells
like Choanoflagellates) (see this helpful movie http://www.youtube.com/watch?v=VLr-_L5HuhU).
The support for the structure (which, like most animals, is largely kept together with the protein
collagen) comes from spicules (based on silica or calcium carbonate). See
http://www.bbc.co.uk/programmes/p010nrzh from Attenborough’s Ark, for some good footage of
sponges and the Venus flower basket sponge.

Nearly all sponges are suspension feeders – they collect food from water passing though
something (a porous body in this case – hence another name for them is Porifera). Flagella of
choanocytes [32.3 and handout] generate water current and the collars trap food, which is then
ingested by phagocytosis. For some nice explanatory animations see
http://www.mesa.edu.au/friends/seashores/sponges1.html

Animal Characteristics

Phagocytosis is one of the main characteristics of animals – ingestion of food (cf. secrete
enzymes to break it down, as in fungi). Another animal characteristic is a layer of cells on the
inside of the organism – endodermis (endo – inside, dermis – skin) [handout; 32.2] formed from a
“pouch” of infolded cells. This layer generates tissue lining the digestive tract in Eumetazoa. The
layer on the outside ecto- or epidermis, generates skin. The pore left where the pouch forms is
called a blastopore, important to remember for later lectures. [32.2 and handout] shows a typical
animal zygote – it undergoes cleavage to generate first a ball then a hollow sphere of cells –
blastula, before the infolding takes place that generates a 2-layered structure.

Eumetazoa fall into two relatively easily distinguishable groups based on body plans: those with
radial symmetry (Radiata) – such as sea anemones and jellies, and those that have only one
plane along which they can be divided into two mirror images – and those that are bilaterally
symmetrical (Bilateria). [32.7] - such as us, lobsters, etc so we will start the next lecture with the
Radiata.

BIOL10511Lecture Key learning outcomes Key words

14 Understand what Protozoa
distinguishes the major Metazoa
groups of animals. Eumetazoa
Understand how single- Radiata
celled animals can Bilateria
collaborate to make a Endodermis
many-celled structure. Blastopore
Understand the main
characteristics of
animals.

Question for (PASS) discussion

The cells that make stalks in Dicty fruiting bodies are genetically distinct from those that make
spores. How/why did this altruistic/suicidal behaviour evolve?

Most sponges are radially symmetrical – why aren’t they in the Radiata?