BIOL 10511 Lecture 7 “Plant Diversity” 1.

Structural and reproductive

advances.

The first aim of this lecture is continued from last time – to continue to show
what important model organisms Chlamy and some of its relatives are. The
handout gives one example of biomedical importance/potential – there are
many others.

Chlamydomonas shares with other green algae and land plants the
possession of chlorophylls a and b with carotenoids as accessory pigments. It
is of historical importance not only as the first genetically engineered
organism (last lecture), but also as the organism that revealed the
phenomenon of cytoplasmic inheritance. In 1954 Ruth Sager crossed two
strains of Chlamydomonas. The cells of one strain were resistant and those of
the other were sensitive to the antibiotic streptomycin. (Streptomycin inhibits
transcription on 70S ribosomes.) The plus strain was resistant, the minus
strain was sensitive. If we understand meiosis in Chlamy (which gives 4
products for each fusion of gametes – two plus and two minus, see last
lecture) in terms of resistance and sensitivity we would expect a one to one
ratio of resistant to sensitive in the progeny.

Sager found a one to one ratio of mating types, but all of the progeny were
resistant to the antibiotic streptomycin (a 4:0 ratio). As we covered in earlier
lectures, genes are carried in chloroplasts and mitochondria (organelles in the
cytoplasm thought to represent the vestiges of engulfed symbiotic organisms)
in addition to those in the nucleus. If the component of plastids or
mitochondria are lost or destroyed from one gamete before or shortly after
fertilisation, as very often happens, their genomes are lost and the offspring
only inherit these genes from one parent (in Chlamy this is the + strain), not
both. So remember that the “genetic make-up” of organisms comprises more
than just the genes in the nucleus.

The second aim of this lecture is to continue to use the green algae to
illustrate the diversity of green photosynthetic organisms, some of which have
fully differentiated sexes and “protect their young”.

Volvox is a hollow sphere, comprising lots of Chlamydomonas-like cells which

remain connected to each other [handout] so form a truly multicellular
organism most correctly referred to as a coenobium (fixed number of cells of
the same species all formed by the same series of cell divisions) – in the
movie I showed and many textbooks it is referred to as a “colony”, but it is one
member of a group that produces organisms with fixed multiples of cells (as
you may have seen in practicals if you are doing them).

For a movie showing how it moves (if you didn’t do the practicals) see
http://www.youtube.com/watch?v=9pjW1cMfTz8&feature=related and for
some nice pictures, including some I showed in the lecture,(but some badly
confused terminology - e.g. should be ‘gonidia’ not gonads! - see
http://www.microscopy-uk.org.uk/mag/indexmag.html?http://www.microscopy-uk.org.uk/mag/artdec03/
volvox.html
Flagella stick out of the extra cellular matrix around the periphery [handout]
and cause the organism to roll through the water. Newly formed coenobia
(clones of the “parent”), produced asexually [handout], are ‘cared for’ inside
the “parent” until they invert – they form a 2-layered structure while they turn
the right way out [handout], using molecules identical to those in many animal
systems during development (of a blastula into a gastrula). Developmental
Biology students are especially recommended to read Nature 424, 499-500
(31 July 2003)”Developmental biology: How to turn inside out”:
http://www.nature.com/nature/journal/v424/n6948/full/424499a.html

Each new clone then ‘hatches’ and swims off to fend for itself [movie]. Volvox
is another “model system” – not only for studies of multicellularity and
developmental biology but also to work out the reasons why reproduction that
includes sex is such a good strategy [see “Sex and stress: linked right from
the start” New Sci 13 August 2005
http://www.newscientist.com/channel/sex/mg18725124.900-sex-and-stress-linked-right-from-
the-start.html

This shows how Volvox “turns sexual reproduction on” in response to heat
stress – formation of molecules that damage DNA causes gamete formation,
so one reason why sexual reproduction may have persisted over time is as a
method of DNA damage repair.

Although Volvox is well differentiated and structurally complex, showing a

division of labour [handout], there are obvious profound limitations on the size
that a hollow sphere of cells can reach. Indeed, Volvox [28.3] is restricted to
relatively undisturbed bodies of water. As in large brown algae, a mass of
intertwined filaments makes a tough and flexible structure and Codium
(common name ‘Dead Man’s Fingers’) is constructed this way [handout]. It
grows in rocky areas subject to considerable wave action, (as you may see if
you go on the BIOL 10622 Majorca field trip). More about Codium in
symbiosis lectures, later in the unit.

Two of the more robust and successful members of the Chlorophyta (green
algae) are Ulva (sea lettuce) a flexible plate of tissue (thallus) or tube
[handout] with a slippery surface – reduces drag [28.21](also see practicals if
you are doing them). Ulva intestinalis is capable of survival in both fresh and
salt water and so can cause ‘fouling’ problems for marine vessels. Ulva
lactuca can also cause huge environmental problems (eg see BBC news 18 th
July http://www.bbc.co.uk/news/world-europe-14182135 and other movies that come
up as related). The poisonous gas reported is probably hydrogen sulphide
gas – the source is not clear, but could be the breakdown produce of a
sulphur- containing osmolyte DMSP, which is abundant in many coastal
marine algae.

Another successful strategy for surviving the rigours of the coastal

environment is secretion of calcium carbonate (as in some stromatolites,
covered earlier). The most famous green alga that does this is Acetabularia –
Mermaid’s Wine Glass [handout]. The calcium carbonate exoskeleton allows
the single cell (up to 10cm long) to survive on very exposed rocky coasts [e.g.
northern coastline of Majorca – BIOL10622]. You may have heard of this
alga before in relation to molecular biology. The central dogma of DNA to
RNA to protein was established in this seaweed [handout]. Because it is
such a big cell it is really easy to manipulate (good for physiologists). There
are different forms, some with fringed caps, others with straight caps, and
transplant experiments between the two allow the identification of messenger
molecules, so it has been important in genetic studies.

A surprisingly successful strategy is adopted by Caulerpa [28.21, handout] –

the whole organism is one huge multinucleate (many nuclei) cell, but it has
remarkable powers of repair, and one form has a really awesome ability to
colonise new environments (and cause considerable environmental
problems). [Access the movie on this - “Movie follow on from lecture 7” in
“additional movies” in BB resources (it is needed for the next eLearning
module, and/or see BSR Caulerpa - “Killer on the loose”].

Reproduction. Some of the Chlorophyta show evidence of evolutionary

“improvements” that are thought to have led to the most successful land
plants. In Chlamydomonas we saw gametes that were identical – termed
isogametes. The green alga Oedogonium (“pond-weed” – in some practicals
you may do) provides yet another example of diversity of structure –
filamentous – but also an evolutionary ‘improvement’ in reproductive terms.
The organism has a high metabolic investment in the female gamete (large
well endowed egg [oocyte - handout] wrapped in a protective wall), but a far
lower metabolic investment in each, much smaller, male gamete. This is
called “oogamous reproduction”. The male gamete is rather like
Chlamydomonas – flagellate (so motile), very little cytoplasm – can now
describe this as a sperm [handout]. The male is strongly attracted to the
female thanks to chemo-attractants [movie], so there is no problem with the
female gamete being non-motile – the female gamete, and therefore the
developing zygote, are housed within the protection of the female parent cell
wall (“parental care”).

It is a very small step from this to the group known as charophytes, which are
probably the ancestral group to all the land plants (handout and more next
time). Coleochaete is the simplest example cited in the textbook [29.3 and
middle of the bottom 3 images on the handout], but Chara (right hand image
at bottom of handout), with its “exoskeleton” of calcium carbonate, is of more
environmental importance so is the only member of this group I expect you to
know about.

BIOL10511Lecture Key learning outcomes Key words

7 Understand importance Coenobium
of green algae eg as Multinucleate
model systems. Exoskeleton
Appreciate structural Oogamy
and reproductive Oocyte (egg)
advances shown by Sperm (male gamete)
green algae. Parental care
Questions for (PASS) discussion

• What are the disadvantages of having a rigid exo-skeleton (plants and

animals)?
• Why can’t we be sure what the earliest land plants looked like?