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37.2 Nonvascular plants are relatively unspecialized, but successful in many

terrestrial environments.
Mosses, Liverworts, and also have specialized food-conducting
cells surrounding those that conduct
Hornworts water.
There are about 24,700 species of Multicellular gametangia are formed
bryophytesmosses, liverworts, and at the tips of the leafy gametophytes
hornwortsthat are simply but highly (figure 37.5). Female gametangia

Seedless vascular plants

Nonvascular plants

Angiosperms
Gymnosperms
adapted to a diversity of terrestrial en- (archegonia) may develop either on the
vironments (even deserts!). Scientists same gametophyte as the male ga-
now agree that bryophytes consist of metangia (antheridia) or on separate
three quite distinct phyla of relatively plants. A single egg is produced in the
unspecialized plants. Their gameto- swollen lower part of an archegonium
phytes are photosynthetic. Sporophytes while numerous sperm are produced in
are attached to the gametophytes and an antheridium. When sperm are re-
depend on them nutritionally to vary- leased from an antheridium, they swim
ing degrees. Bryophytes, like ferns and with the aid of flagella through a film of
certain other vascular plants, require dew or rainwater to the archegonia.
water (for example, rainwater) to re- One sperm (which is haploid) unites
produce sexually. It is not surprising with an egg (also haploid), forming a
that they are especially common in diploid zygote. The zygote divides by
moist places, both in the tropics and tem- mitosis and develops into the sporo-
perate regions. phyte, a slender, basal stalk with a swollen
Most bryophytes are small; few exceed 7 capsule, the sporangium, at its tip. As the
centimeters in height. The gametophytes sporophyte develops, its base is embedded
are more conspicuous than the sporo- in gametophyte tissue, its nutritional
phytes. Some of the sporophytes are com- source. The sporangium is often cylindrical
pletely enclosed within gametophyte tissue; or club-shaped. Spore mother cells within
others are not and usually turn brownish or the sporangium undergo meiosis, each pro-
straw-colored at maturity. ducing four haploid spores. In many mosses
at maturity, the top of the sporangium pops
off, and the spores are released. A spore
Mosses (Bryophyta)
that lands in a suitable damp location may
The gametophytes of mosses typically germinate and grow into a threadlike struc-
consist of small leaflike structures (not ture that branches to form rhizoids and
true leaves which contain vascular tissue) buds that grow upright. Each bud devel-
arranged spirally or alternately around a ops into a new gametophyte plant consist-
stemlike axis (figure 37.4); the axis is an- ing of a leafy axis.
chored to its substrate by means of FIGURE 37.4 In the Arctic and the Antarctic, mosses
A hair-cup moss, Polytrichum
rhizoids. Each rhizoid consists of several are the most abundant plants, boasting not
(phylum Bryophyta). The
cells that absorb water, but nothing like leaves belong to the only the largest number of individuals in
the volume of water absorbed by a vascu- gametophyte. Each of the these harsh regions, but also the largest
lar plant root. Moss leaves have little in yellowish-brown stalks, with the number of species. Many mosses are able to
common with true leaves, except for the capsule, or sporangium, at its withstand prolonged periods of drought, al-
superficial appearance of the green, flat- summit, is a sporophyte. though they are not common in deserts.
tened blade and slightly thickened midrib Most are remarkably sensitive to air pollu-
that runs lengthwise down the middle. tion and are rarely found in abundance in
They are only one cell thick (except at the midrib), lack or near cities or other areas with high levels of air pollu-
vascular strands and stomata, and all the cells are haploid. tion. Some mosses, such as the peat mosses (Sphagnum),
Water may rise up a strand of specialized cells in the can absorb up to 25 times their weight in water and are
center of a moss gametophyte axis, but most water used by valuable commercially as a soil conditioner, or as a fuel
the plant travels up the outside of the plant. Some mosses when dry.

738 Part X Plant Form and Function

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Liverworts (Hepaticophyta) Sperm

The old English word wyrt means Antheridium
Zygote
plant or herb. Some common liver-
worts have flattened gametophytes with FERTILIZATION Developing
sporophyte in
lobes resembling those of liverhence archegonium
the combination liverwort. Although
the lobed liverworts are the best-known
representatives of this phylum, they Egg
Mature
constitute only about 20% of the sporophyte
Archegonium
species (figure 37.6). The other 80%
are leafy and superficially resemble
mosses. Gametophytes are prostrate in-
stead of erect, and the rhizoids are one- n 2n
celled.
Some liverworts have air chambers
containing upright, branching rows of
photosynthetic cells, each chamber Male Female Sporangium
having a pore at the top to facilitate gas
exchange. Unlike stomata, the pores
Gametophytes
are fixed open and cannot close. Parent
Sexual reproduction in liverworts is gametophyte
MEIOSIS
similar to that in mosses. Lobed liver-
worts may form gametangia in um- Bud Mitosis
brella-like structures. Asexual repro- Rhizoid
duction occurs when lens-shaped Germinating
Spores
pieces of tissue that are released from spores
the gametophyte grow to form new
FIGURE 37.5
gametophytes. Life cycle of a typical moss. The majority of the life cycle of a moss is in the haploid
state. The leafy gametophyte is photosynthetic, while the smaller sporophyte is not, and
Hornworts (Anthocerotophyta) is nutritionally dependent on the gametophyte. Water is required to carry sperm to
the egg.
The origin of hornworts are a puzzle.
They are most likely among the earliest
land plants, yet the earliest hornwort
fossil spores date from the Cretaceous
period, 65 to 145 million years ago,
when angiosperms were emerging.
The small hornwort sporophytes re-
semble tiny green broom handles rising
from filmy gametophytes usually less
than 2 centimeters in diameter (figure
37.7). The sporophyte base is embed-
ded in gametophyte tissue, from which
it derives some of its nutrition. How-
ever, the sporophyte has stomata, is
photosynthetic, and provides much of
the energy needed for growth and re-
production. Hornwort cells usually
have a single chloroplast.
FIGURE 37.6 FIGURE 37.7
A common liverwort, Marchantia Hornworts (phylum Anthocerotophyta).
The three major phyla of
(phylum Hepaticophyta). The Hornwort sporophytes are seen in this
nonvascular plants are all relatively
sporophytes are borne within the tissues of photo. Unlike the sporophytes of other
unspecialized, but well suited for
the umbrella-shaped structures that arise bryophytes, most hornwort sporophytes are
diverse terrestrial environments.
from the surface of the flat, green, creeping photosynthetic.
gametophyte.

Chapter 37 Evolutionary History of Plants 739