Lecture 11 Notes 2013

Lecture 11 Plant diversity 5. From spores to seeds.
Case study of water

ferns.
As we shall see later in the unit, many organisms that arose on dry land have
evolved into aquatic inhabitants, and one unusual group of ferns lives out its
life in freshwater habitats – the water ferns. The first aim of this lecture is to
review members of this important group.
Salvinia molesta (Kariba weed) has a floating rhizome with leaves that
protrude out of the water [handout] bearing waxy “egg-beater” hairs that help
make it water repellent. It can double its dry weight in 2½ days. The asexual
spread of this fern has caused enormous environmental problems in
developing countries, and also where introduced as an ornamental (cf.
Caulerpa) e.g. USA (see, for example, http://www.youtube.com/watch?v=S9Mf95GDGjgtp;
http://www.youtube.com/watch?v=xSOIfFnradw ) and Australia. Biological control can be
achieved using a species of weevil that would rather die than eat anything
else [Nature 294, 5 November 1981; and BSR article in BB resources]. See
also http://www.csiro.au/Outcomes/Safeguarding-Australia/salvinia-
control.aspx for some interesting history of this weed and biological control).
Another tropical water fern of use to man as an ornamental is Azolla. It is now

causing some concern in the UK as growth used to be killed off in winter but
now survives, and can clog water-ways [see handout], but it is mainly
beneficial, being of enormous importance in rice paddies as “green” fertiliser
[also mentioned in lecture 2]. It has a cyanobacterium – Anabaena azollae,
living inside its leaves [handout] (which, like free-living cyanobacteria, can
contribute to the nitrogenous needs of rice – when the rice-growing season
reaches its peak, the heat and dryness kills the fern and the fixed nitrogen
becomes available to the rice) [more about this in later lectures on organisms
in partnerships].
These water ferns owe their success primarily to asexual reproduction –

rhizomes with horizontal growth and branching (cf. Caulerpa; Equisetum;
bracken) – but they are heterosporous, like Selaginella (last lecture) - a habit
that started (probably in several groups of plants) in the Carboniferous era
[handout] and generated the most successful group of all extant plants – seed
plants. The main aims of the rest of this lecture are 1. to review diversity of
gymnosperms – the smallest of the two groups of extant seed plants by far
but encompassing an enormous diversity of vegetative form, environmentally
and economically important plants; 2. to show how spore-bearing plants
changed into seed-bearers.
Free-living gametophytes such as those of most ferns are relatively vulnerable

in comparison with sporophytes – they have no vascular tissue and negligible
cuticle – vulnerable. The heterosporous pterophytes such as Selaginella and
some Carboniferous lycophytes (not well covered in Campbell text book so
see the Ennos and Sheffield book p108-110 if you want more detail) have
sporangia of 2 different types. Megasporangia produce megaspores – large
spores with metabolic reserves to provide for the growing embryo.
Microsporangia produce small spores, which generate only numerous small
motile sperm. These plants have no free-living photosynthetic gametophyte
plant – fertilisation is via motile sperm, but this event happens within the
protection of the megaspore wall - “internal fertilisation”, in addition to the
“parental care” aspect of the female gamete being protected on the parent
organism (we will see a similar story of improvements during evolution in the
animal groups in later lectures).
The evolutionary trends we have considered so far have led plants to a higher
and higher investment in the female line, and continual reduction in the
haploid (gametophyte) stage of the life cycle.
Seed plants – if instead of dispersing their spores, Selaginella or

Carboniferous lycopods retained their megaspores on the parent plant (this
certainly happens/happened in some species) – the new generation would be
well protected and provided for by the parent sporophyte, with the very much
reduced (non-photosynthetic) gametophyte completely dependent upon it, but
protected. This is how we think seed plants arose.
Handout [30.3] shows how the megasporangium of pterophytes may have

been reduced to contain only a single megaspore, surrounded by protective
tissue from the parent plant – now we call this structure an ovule. This would
have been fertilised on the parent plant by a sperm generated by a pollen
grain (microspore). The seed (with fertilisation already safely done, and
embryo formed) provides another opportunity for the plant to disperse its
genes (in addition to that provided by its pollen grains). The seed has a coat
to protect it, a food supply for the embryo and, very often, the means to
ensure efficient travel far from parent plant (e.g. wing-like structures, etc,
more about this in later lectures).
The first successful organisms with seeds were gymnosperms [see handout
and 30.5] (“naked seed” plants) in which 4 phyla of extant plants are
recognised. Ginkgophyta – only living representative is Ginkgo biloba
[handout; 30.5; leaf in the lecture and see Field Specimen # 2 in the front
quad of Stopford]. This taxon was very much more successful in earlier times
– now it is a living fossil (probably extinct in the wild) that has survived
because of its huge environmental tolerance, horticultural and economic
importance. It can tolerate extremes of temperature and desiccation, and
even atmospheric pollution – hence it is now planted in cities all over the
world.
Ginkgo is a deciduous plant – losing leaves is one strategy for surviving long
periods of low temperature and/or light [watch Field Specimen #2 over the
next few weeks .Also see BSR Maidenhair tree article in resources, which
outlines some of the horticultural use and claimed health benefits of Ginkgo-
based products NB since this was published there have been many more
publications but also warnings about the biomedical hazards of some Ginkgo
products; also see handout].
Ginkgo generates pollen which travels through the air to the female parts
[handout] and then generates motile sperm, which are large and not
streamlined like those of the ferns and bryophytes – but then they don’t have
to swim very far. Internal fertilisation leads to the formation of seeds with (very
smelly!) fleshy exterior, which probably once appealed to an animal that would
have helped to disperse the seeds (but which has become extinct – most
animals now avoid the fleshy coatings on the seeds).
[watch the sequence in the following – which starts with foliage unfurling in
time lapse to give an idea of how attractive the new foliage looks in spring,
then shows sperm on its own, then inside an archegonium, then shows sperm
of a cycad [see later in this lecture], then goes into a sequence on the cellular
slime mould Dictyostelium, which will be relevant to lecture 16 in this series]
http://www.youtube.com/watch?v=9OVeE-28RyA
Cycads were very important in the Mesozoic era (“Age of cycads” to plant
scientists – “age of dinosaurs” to zoologists – 250 mya). They have a very
limited present day distribution but are still important horticulturally. The
trunks of arborescent forms look like those of tree ferns – these too have very
limited woody vascular tissue [handout; 30.5], and are mainly supported by
large woody leaf bases and the vascular traces that lead into them. The
leaves are “evergreen” - very big and metabolically expensive to make, so the
plants generate leaves only every few years, even in the best conditions, and
are very slow growing. Very infrequently, instead of producing leaves, the
plants will produce cones – very large woody structures – see time lapse
taken over a 10 day period http://www.youtube.com/watch?v=Ryk6PEmDzfo&NR=1.
The years when this happens, the plant does not have sufficient reserves to
generate leaves.
Cycads are dioecious – di = two, oikos = house, i.e. separate male and
female plants. Male plants therefore have to transmit gametes to females.
This is done with pollen grains. It was once thought that their pollen grains
were carried by the wind (a common misconception is that only flowering
plants employ animal vectors) – but pollination is now known to be insect-
mediated in many species. In some, the cones heat up, emit volatile odours,
particular insects are attracted to feed or mate on the cones. In others there is
a push-pull system of attracting and repelling thrips [see handout and “Ancient
plant has hot stinky sex” New Sci 4 th October 2007
http://www.newscientist.com/article/dn12739-ancient-plant-has-hot-stinky-
sex.html (includes YouTube movie). Whatever the mechanism, insects help in
pollination by transporting the pollen directly to where it needs to be. Once
the pollen is inside the female cone, germination right inside the cone again
generates motile sperm.
BIOL10511Lecture Key learning outcomes Key words

11 Understand success Internal fertilization
(mostly asexual Ovule
reproduction) of water Seed
ferns. Deciduous
Understand how spore- “Evergreen”
bearing plants gave rise Dioecious
to seed-bearers.
Questions for (PASS) discussion
• Are there any advantages to having individuals of separate sexes (as

opposed to hermaphrodites – monoecious plants).
• If you were a plant would you be deciduous or evergreen? (can you
think of mammalian equivalents of these 2 different strategies for over-
wintering?)

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Lecture 11 Plant diversity 5. From spores to seeds.

Case study of water

Another tropical water fern of use to man as an ornamental is Azolla. It is now

These water ferns owe their success primarily to asexual reproduction –

Free-living gametophytes such as those of most ferns are relatively vulnerable

Seed plants – if instead of dispersing their spores, Selaginella or

Handout [30.3] shows how the megasporangium of pterophytes may have

BIOL10511Lecture Key learning outcomes Key words

• Are there any advantages to having individuals of separate sexes (as

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