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It is well established that living things can trace their ancestry to early life forms that were
restricted to aquatic environments. Because of lifes origin in the sea, scientific discussions of
the evolution of plants (and animals) often emphasize key evolutionary novelties that allowed
organisms to invade terrestrial (land-based) environments. In this lab, you will compare two
groups of plants - the Bryophytes and the Ferns/Fern Allies with respect to key evolutionary
features that enabled plants expansion into terrestrial habitats.
Bryophytes lack many of the structural adaptations typical of higher plants, including true
roots, stems, and leaves. Despite their simplicity, bryophytes have become quite successful in a
variety of terrestrial environments. Ferns and fern allies are especially significant because they
represent the earliest group of plants to have vascular tissues (xylem and phloem) which
transport water, minerals, and sugars throughout the plant body. Because of this advance, ferns
and fern allies are able to achieve much larger body sizes and occupy a broader diversity of
habitats than their bryophyte predecessors.
ALTERNATION OF GENERATIONS
Well assume that by now, youve all mastered the story of your own creation: two haploid
(haploid, or 1n = 1 set of chromosomes) gametes, one sperm and one egg, fused to form a diploid
(diploid, or 2n = 2 sets of chromosomes) zygote (fertilized egg) that divided repeatedly by
mitosis to form a mass of cells known as an embryo. The embryo continues to grow and
differentiate until it reaches maturity as an adult. At some point in your life cycle, you (the
adult) produce diploid sex cells in your sex organs that will divide by meiosis to form haploid
gametes (sperm or eggs). And, the cycle is perpetuated. Sound familiar?
It is helpful in our understanding of plant reproduction if we draw upon what we already know
about our own reproduction and seek the similarities. Of course, we would be negligent if we
didnt also address the differences there are more than a few! Perhaps the most striking
difference between plant and animal life cycles is that plants spend part of their life cycle in a
haploid (1n) phase, and part of their life cycle in a diploid (2n) phase. This type of life cycle,
known as alternation of generations, is characteristic of all plants, and also occurs in a few
other non-plant organisms, including most algae. You, as well as most animals, only exist in a
diploid (2n) state regardless of where you are in your life cycle.
With the exception of the bryophytes, most organisms that you recognize as plants represent
only the diploid sporophyte generation of that plant. Sporophyte literally means spore plant
or spore-producing plant. Sporangia (singular: sporangium) are the structures located on the
body of the sporophyte where spores are produced. Diploid cells inside sporangia undergo
meiosis to produce haploid spores. The haploid spores germinate (grow) to give rise to the other
generation: the haploid gametophyte.
Gametophyte literally means gamete plant or gamete-producing plant. Not surprisingly
then, you will find structures on the gametophyte where the gametes (sperm and eggs) are
produced. Antheridia (singular: antheridium) are the structures where sperm are produced;
archegonia (singular: archegonium) are structures where eggs are produced. Through a variety
of sometimes remarkable, if not downright bizarre mechanisms, sperm and egg are eventually
united and fuse to form the diploid zygote that will eventually grow by mitosis into a new
sporophyte.
Sperm
fuse to form
cells in ovaries
undergo meiosis
Female
Zygote
Eggs
growth
by
mitosis
Embryo
growth by mitosis
growth by
mitosis
Haploid
Sporophyte (2n)
Gametophyte (1n)
Diploid
growth
by mitosis
produces
Zygote (2n)
Gametes (1n)
fuse to form
Angiosperms
Gymnosperms
Ferns
flowers, fruits
seeds
*Bryophytes
pollen
vascular tissue
Green Algae
dominant sporophyte
dominant gametophyte
flagellated sperm
Chlorophylls a, b
The Phylum Bryophyta includes 3 groups of plants the mosses, liverworts, and hornworts.
Bryophytes are probably the best example of what the earliest plants may have looked like. The
body of bryophytes is relatively undifferentiated and lacks true roots, stems, and leaves. The
bryophyte body is sometimes referred to as "thallus", a term used to refer to generalized,
undifferentiated plant tissue. Threadlike structures called rhizoids can be found on the lower
surface of the thallus that are considered important for anchoring the plant to the substrate
but do not have a role in water absorption.
Bryophyte distribution tends to be somewhat restricted to moist environments. Bryophytes
lack vascular tissues (xylem and phloem) and are therefore unable to transport water and
dissolved substances from one part of the plant body to another. Because of their reliance on
diffusion for transport, bryophytes cannot obtain large body sizes that might be typical of
higher plants. Water is also required for bryophyte reproduction - flagellated sperm released
into the environment must swim through a liquid medium in order to make their way to an egg in
a nearby archegonium. Because of these constraints, bryophytes are found predominantly in
moist areas in forests and close to streams, bogs, wetlands, ponds, etc. . . Having said that,
some bryophytes have an also be found in rather extreme environments, such as deserts, the
Arctic and Antarctic, and mountaintops well above timberline.
Ecologically, bryophytes are important for a number of reasons. In some habitats, they are the
primary sinks for global atmospheric carbon, fixing vast quantities of carbon into biologically
usable carbohydrates. Bryophytes are also important early colonizers of barren landscapes such
as bare rock, mine tailings, and volcanic fields. Like lichens (a symbiotic relationship between
algae and fungi), many bryophytes are considered important indicator species because of
their extreme sensitivity to air, water, and soil pollution.
*Ferns
flowers, fruits
seeds
Bryophytes
pollen
vascular tissue
Green Algae
dominant sporophyte
dominant gametophyte
flagellated sperm
Chlorophylls a, b
The ferns and ferm allies represent the earliest group of plants in which vascular tissues are
present. Vascular plants are categorized into 2 groups based on whether or not they are capable
of producing seeds - the seedless vascular plants and the seed plants. The seedless vascular
plants (also known as the ferns and fern allies) are represented by four phyla: Psilotophyta,
Lycophyta and Sphenophyta (the fern allies) and Pterophyta (the true ferns). The primary
difference between the true ferns and their allies is the presence of large, complex leaves
called megaphylls found in the ferns. The individual phyla are described briefly below.
If available, examine a specimen of Psilotum for this group. The plant body consists entirely of
dichotomously-branched (forked) stems, as this group lacks true roots and leaves. The scalelike appendages on the branches are sometimes referred to as prophylls (pro = first or
before; phyll = leaf), but are not considered true leaves because the vascular tissues do not
extend into them. The belowground portion of the plant also consists entirely of a modified
stem, or rhizome, with tiny absorptive hairs called rhizoids. Sporangia are borne in groups of 3
at the tips of short side branches. Psilotum produces only 1 kind of spore, hence is termed
homosporous (meaning same spore). The spore germinates underground and gives rise to a
subterranean, bisexual gametophyte, bearing both the male antheridia and the female
archegonia.
Examples of lycophytes include members of the genera: Selaginella, Lycopodium, and Isoetes.
Lycophytes have true roots, stems, and leaves, though the arrangement of the parts can vary
dramatically from one species to the next. The leaves of this group are known as microphylls
(meaning small leaves) as they generally contain just a single trace of vascular tissue to serve
the whole leaf. The sporangia of lycophytes are borne at the base of leaves called sporophylls
(meaning spore leaves) which, in some cases, are arranged into structures that resemble
cones, called strobili (singular: strobilus). The sporophylls are not fundamentally different from
any of the other leaves on the plant, except that they happen to support sporangia, thus earning
them a distinguishing title.
All the living sphenophytes are currently represented by a single genus: Equisetum. Equisetum is
easily identified by its characteristic jointed stems with whorls of leaves at the nodes. Roll the
stem of Equisetum in your fingers to feel the ribs another characteristic feature of this
plant. Whereas the sporangia of lycophytes are borne on leaves, the sporangia of sphenophytes
are borne on stems (as in the psilophytes). However, unlike the psilophytes, those modified
stems bearing the sporangia are arranged into strobili that somewhat resemble the strobili of
lycophytes.
The key characteristic separating the fern from their allies, is in their possession of large,
complex leaves known as megaphylls (meaning large leaves). Apart from size, megaphylls
differ from microphylls in containing more than one vein (bundle of vascular tissue). From ferns
through the flowering plants, leaf architecture is one of the primary ways botanists describe
differences in morphology among species. The terms below apply to the morphology of fern
leaves, in particular. Be sure that you can relate the terms to the actual structures on the fern
specimens provided.
frond = leaf of fern, including the petiole
petiole = the stalk-like structure of the leaf which connects it to the stem
lamina (also, blade) = the broad part of the leaf, often flattened
pinnae = leaflets, or divisions of the lamina
rachis = extension of the petiole to which the pinnae are attached
Most, but not all, ferns bear sporangia on their leaves in clusters called sori. Some species
produce separate vegetative (non-reproductive) and fertile fronds. In other species, the fronds
are all equivalent and sori may be found on any of them. The location of the sori varies from
species to species check the specimens in lab for sori on the underside of leaves or at the leaf
margins. Do any of the specimens have separate fertile and vegetative leaves? Do any of the
specimens produce indusia - protective caps that enclose the sori?
The life cycle of the ferns and allies is based on the same model described for bryophytes, but
not surprisingly, some of the structures look different or are named differently. As mentioned
in our study of bryophytes, the bryophytes are the only plant group in which the gametophyte is
the dominant generation. Hence, the plant bodies you recognize as a fern or fern ally
represent the dominant sporophyte generation of the plant. It is unlikely that you will ever
encounter the discrete gametophytes (called prothalli, singular prothallium) on a casual walk
through the woods as they tend to be very small and some develop completely belowground.
However, like the bryophytes, this group has flagellated sperm produced in antheridia that
require water in order to swim to eggs and complete the sexual life cycle. During its early
development, the young sporophyte is nutritionally dependent on the gametophyte, but soon
becomes independent, acquiring all its nutrition via photosynthesis.
ACTIVITIES:
1. Examine the liverwort specimen provided (probably Marchantia). In Marchantia, the sexes are
separate. In the male gametophyte, stalked, disk-like structures that contain sperm-producing
antheridia are called antheridiophores. In the female gametophyte, archegoniophores are
the stalked, umbrella-like structures where egg-producing archegonia can be found.
Draw and label the thallus, gemmae cups, rhizoids, and antheridia or archegonia (if present) of
the gametophyte.
Look for sporophytes. If present, these will be located on the underside of the
archegoniophore. Draw and label the foot, seta, and sporangium of the sporophyte containing
spores.
For each of your drawings, indicate whether each structure you labeled is 2n or 1n.
2. Repeat #1, but for one of the mosses provided. Draw and label the foot, seta and sporangium
of a sporophyte and the thallus, rhizoids, and antheridia/archegonia of the gametophyte. (For
many mosses, it may be very difficult to distinguish the antheridia from archegonia - especially
if sporophytes are absent). As with the liverwort, represent both the gametophyte and
sporophyte generation and indicate whether each structure you label is 2n or 1n.
3. Examine a fern prothallium from your terrarium under a dissecting scope, then make a wet
mount of it. Draw and label the rhizoids, antheridia, and archegonia.
4. Examine the array of ferns and fern allies on display, including prepared slides and live or
preserved tissues available for you to dissect or make wet mounts of (check with you TA before
dismembering specimens!).
Draw an example of both a fern and one fern ally (sporophyte phase), labelling all relevant
structures (which structures you label will depend on which specimens you choose). Make sure
you can identify key structures, including roots, stems, leaves, rhizomes, rhizoids, microphylls,
megaphylls, sporangia, strobili, sori, fronds (including lamina, pinnae, petiole and rachis),
gametophytes, sporophytes, etc. . . Compare your 2 drawings noting similarities and differences.
Feel free to consult lab manuals, posters, and texts provided in lab (or websites) to help you in
identifying structures.
POSTLAB QUESTIONS:
1. Bryophytes lack the vascular (conducting) tissues found in all higher plants. Given this, how
do water, minerals, and sugars get dispersed throughout all the cells of the plant body? How
does this constrain the body size achievable by a bryophyte?
3. a) Is the sporophyte of a fern capable of living independently? Where does its nourishment
(food) come from?
b) Is the fern gametophyte capable of living independently? Where does its nourishment (food)
come from?
4. Bryophytes are considered the most primitive of the plants, but represent a significant
advance over their evolutionary precursors the green algae. What features of bryophytes
continue to restrict their distribution on land?
5. What evolutionary adaptations in the ferns/allies distinguish them as being more advanced
than their bryophyte predecessors and account for their success and expansion in terrestrial
habitats? Are there any features of ferns that continue to restrict their distribution?
Images: Bryophytes.
http://www.ksu.edu/organismic/bryophytes.htm#
http://www.ksu.edu/organismic/bryophytes.htm#
Images: Ferns.
http://web.vet.cornell.edu/CVM/HANDOUTS/plants/Pter
idium%20drawing.jpg
http://www.hcs.ohio-state.edu/hcs300/svp2.htm
http://www.extension.umn.edu/specializations/enviro
nment/images/lycopodiumdraw.gif
http://scitec.uwichill.edu.bb/bcs/bl14apl/pteridoequisetum.jpg
http://www.ujfgrenoble.fr/JAL/Choler/BE
V/prat/demo/pter/psilotum.JPG