BIOL 10511 Lecture 8 Plant diversity 2.

Bryophytes (“land plants”)

Charophytes such as Chara have eggs housed within a protective layer of cells
[handout] – “parental care”. However, although the plants have an exoskeleton of
calcium carbonate, they cannot prevent themselves from drying out and dying when
exposed to the air for long periods. Fossil charophyte embryos bear striations
indicating where the surrounding protective cells were when the plant was alive,
even after hundreds of thousands of years [handout]. One reason for this is a highly
resistant polymer that surrounds the embryo: sporopollenin (the same as you find
around pollen grains). Biochemical advances produced a range of such tough
materials in the ancestors of land plants – in Chara, which grows in shallow water,
sporopollenin prevents zygotes drying out and dying if the waterway dries up. The
aim of this lecture is to explore the evolutionary steps like this that eventually led to
the colonisation of land. [see Chapter 5 in Ennos and Sheffield for further reading on
this, and many of the images used to illustrate this lecture]

Alternation of generations. In Chara, the zygote undergoes meiosis to produce

haploid spores, as in Chlamydomonas. Not all algae are like this, but as
charophytes are thought to have given rise to land plants we assume that mutation
resulted in delay in meiosis [handout; 13.6b]. Gametangia is the word for sex organs
(which produce gametes) and the word archegonium is the plant science term
preferred to oogonium for the cellular structure that houses the egg cell (oocyte). An
archegonium has at least one layer of cells that protects and nourishes the first
stages of the new generation (cf. Oedogonium, where the egg was simply housed
within the cell wall of the original cell). Mitotic divisions in the embryo generates a
multicellular body before meiosis, so you get a structure that can generate more
spores per zygote. (Sexual reproduction is usually more expensive metabolically
and riskier than asexual reproduction, so it pays an organism to maximise its output
of products). We can now call the plant that produces the gametes a gametophyte,
and the structure that generates spores, a sporophyte (spore-producing plant).

Bryophytes (bryo = moss, phyton = plant) are organisms that have multicellular
sporophytes that remain attached to the parental gamete-producing plant.

Structural diversity of vegetative form.

There are three groups of bryophytes [29.9], all of which are considered to be “non-
vascular” plants. The earliest forms (see handout) were liverworts (e.g.29.7 [handout
from last time]; the simplest of which are flat plates of tissue – thalloid. The additional step
essential for this to allow them to survive the terrestrial
environment was the development of a cuticle (lipid
and hydrocarbon polymers impregnated with wax,
making a layer that helps to prevent water loss).
Those more than a few cells thick would have
encountered gas exchange problems – pores in the
top surface allow CO2 in and O2 out but as they can’t
be opened and closed (cf. stomata) liverworts are
prone to desiccation and only found in damp places
(such as the corner of the front quad in Stopford).
There is a huge diversity of liverworts, especially in
tropical regions, many of which are divided into lobes - called “leafy” liverworts (not
true leaves - just expansions of the surface area that make them somewhat more
efficient at catching the sun than the thalloid types but which are only a single cell
thick).

Thalloid and leafy liverworts [29.9 and handout] number at least 9,000 species.
Hornworts are like a thalloid liverwort but with horn-like sporophytes – a tiny group of
about 100 spp – no need to learn. The most familiar and successful group, mosses,
number >15,000 species. Mosses are an environmentally important group - found
on every continent in the world, including Antarctica – tolerant of very low and quite
high temperatures and some can regenerate after weeks or even centuries of low
temperatures or desiccation [handout]. They only tolerate low temperature or
desiccation, they don’t continue to metabolise. The day to day on/off existence of
many mosses is tied in with environmental humidity [movie] – leaves tolerate losing
almost all their water content, many can curl round stems to expose dead tissue,
therefore preserving moisture; uncurl and carry on metabolising when re-hydrated. A
good example of a local species that can do this is Polytrichum – Bank hair moss
(more next lecture on this).

Reproductive diversity.

All liverworts produce the same sort of sporophyte – small structures raised above
the parent gametophyte, non-photosynthetic and completely dependent on the
parent gametophyte for nutrition [handout]. Sporangia (the spore-containing
structures at the top) contain two types of cells – elaters and spores [handout].
Sporangia split open when dehydrated and the elaters (programmed cell death –
apoptosis, generates these thickened, specialised cells) react to changes in water
content (hygroscopic movements) and help fling spores into the air [handout].

Sexual reproduction relies on motile male gametes (from male gametangia called
antheridia), and female gametes housed in archegonia, just as in charophytes, but
sometimes there is not a film of water between the male and female plants.
Conocephalulm conicum is one liverwort that has a brilliant way to get around this
problem – read “Airborne sperm of Conocephalum conicum” Journal of Plant
Research 121 68-71 2008 10.1007/s10265-007-0128-6 and scroll down to the
electronic supplemental material to watch the movie.

Moss sexual reproduction is similar to liverworts and charophytes [29.8; movie] –

tiny, streamlined, spirally-shaped motile sperm are produced in vast numbers, which
until relatively recently were assumed always to rely on a film of water or rain splash
to reach female structures/plants but now we know that small arthropods can be
important too [if interested in plant-animal interactions see “Microarthropods mediate
sperm transfer in mosses” Science 1 Sept. 2006
http://www.sciencemag.org/content/313/5791/1255.full]

Female mosses have large, non-motile, well-endowed eggs, protected on the parent
plant – the egg cells are down at the bottom of a long-necked archegonium, down
which only the fittest sperm can swim to fertilise the eggs (the sperm shed most of
their cytoplasm, becoming streamlined and corkscrew-shaped, which helps them
negotiate the tortuous pathway down to the egg [movie]).

Moss sporangia are very different from those of liverworts. They do not have elaters,
but most are otherwise similar as they act via hygroscopic movements [movie]. The
handout shows some of the range of moss peristomes (“around the rim” – structures
 that cap off the sporangia and allow the control of spore release). Some act like little
pepper pots, opening and closing in response to dry and wet air respectively, others
have teeth that bend right back and fling the spores out [handout]. This can
generate up to 70 million spores per sporangium (i.e. loads more meiotic products
than Chlamy or Chara can generate when meiosis is the next step after formation of
a zygote), and a mechanism to make sure they are not all released at the same time
(The textbook is over-simple; movie and handout give some idea of huge diversity).

One exception to the use of hygroscopic movements to aid spore dispersal is shown
by the moss Splachnum (drawing G on the handout). This genus has species which
release odours that resemble carrion, or dung, and so attract carrion/dung flies. The
sticky spores on the central protrusion of the sporangium get stuck to the feet of the
flies, and so get distributed to their preferred habitat when the flies next feed.

BIOL10511Lecture Key learning outcomes Key words

Understand importance Bryophyte
8 and diversity of Gametangium
bryophytes. Archegonium
Antheridium
Understand alternation Sporophyte
of generations. Gametophyte
Liverwort
Elater
Peristome
Hygroscopic
Moss
Sporangium

Questions for (PASS) discussion.

• Given that the earliest land plants are thought to have been bryophytes, what
kind of animals might have helped disperse their sperm or spores?
• What are the advantages and disadvantages of relying on animal vectors to
distribute propagules?