Seedless Plants

What types of
plants are
seedless?
• Ferns, horsetails and
mossess
• Found In cool, moist,
shaded areas in the
forest
• Usually grow near the
ground
• Reproduce sexually by
spores
• Monophyletic
Early Plant Life

The Plant Kingdom

• There are more than 300,000 species of catalogued plants
and 260,000 are plants that produce seeds.
• Mosses, ferns, conifers, and flowering plants are all members of the
plant kingdom.
• Most biologists also consider green algae to be plants, although
others exclude all algae from the plant kingdom.
ALGAE AND EVOLUTIONARY PATHS TO
PHOTOSYNTHESIS
• Some scientist exclude algae in plant kingdom but the other
consider it to be plants.
• All algae are photosynthetic—that is, they contain some form of a
chloroplast—they didn’t all become photosynthetic via the same
path.
• The ancestors to the green algae became photosynthetic by
endosymbiosing a green, photosynthetic bacterium about 1.65
billion years ago.
• That algal line evolved into the Charophytes, and eventually into the
modern mosses, ferns, gymnosperms, and angiosperms. Their
evolutionary trajectory was relatively straight and monophyletic. In
contrast, the other algae—red, brown, golden, stramenopiles, and so
on—all became photosynthetic by secondary, or even tertiary,
endosymbiotic events; that is, they endosymbiosed cells that had
already endosymbiosed a cyanobacterium.
• The different views on whether all algae are Plantae arise from how
these evolutionary paths are viewed. Scientists who solely track
evolutionary straight lines (that is, monophyly), consider only the
Charophytes as plants. To biologists who cast a broad net over living
things that share a common characteristic (in this case,
photosynthetic eukaryotes), all algae are plants.
Plant Adaptations to Life on Land
• Water has been described as “the stuff of life.”
• The cell’s interior is a watery soup: most small molecules dissolve
and diffuse, and the majority of the chemical reactions of
metabolism take place.
• Desiccation, or drying out, is a constant danger for an organism
exposed to air.
• Water also provides buoyancy to organisms.
• On land, plants need to develop structural support in a medium that
does not give the same lift as water. The organism is also subject to
bombardment by mutagenic radiation, because air does not filter
out ultraviolet rays of sunlight.
• Additionally, the male gametes must reach the female gametes
using new strategies, because swimming is no longer possible.
To balance these survival challenges, life on land offers several
advantages.
• First, sunlight is abundant. Water acts as a filter, altering the
spectral quality of light absorbed by the photosynthetic pigment
chlorophyll.
• Second, carbon dioxide is more readily available in air than in
water, since it diffuses faster in air.
• Third, land plants evolved before land animals; therefore, until dry
land was colonized by animals, no predators threatened plant life.

• Tolerance. Many mosses, for example, can dry out to a brown and
brittle mat, but as soon as rain or a flood makes water available,
mosses will absorb it and are restored to their healthy green
appearance.
• Colonize environments with high humidity, where droughts are
uncommon. Ferns, which are considered an early lineage of plants,
thrive in damp and cool places such as the understory of temperate
forests.
• The most successful adaptation solution was the development of
new structures that gave plants the advantage when colonizing new
and dry environments.
• Four major adaptations are found in all terrestrial plants:
• The alternation of generations, a sporangium in which the spores
are formed, a gametangium that produces haploid cells, and apical
meristem tissue in roots and shoots. The evolution of a waxy cuticle
and a cell wall with lignin also contributed to the success of land
plants. These adaptations are noticeably lacking in the closely
related green algae—another reason for the debate over their
placement in the plant kingdom.
Alternation of Generations
Alternation of generations describes a life cycle in which an organism
has both haploid and diploid multicellular stages
• Haplontic refers to a lifecycle in which there is a dominant haploid
stage, and diplontic refers to a lifecycle in which the diploid is the
dominant life stage. Humans are diplontic. Most plants exhibit
alternation of generations, which is described
as haplodiplodontic: the haploid multicellular form, known as a
gametophyte, is followed in the development sequence by a
multicellular diploid organism: the sporophyte. The gametophyte
gives rise to the gametes (reproductive cells) by mitosis. This can be
the most obvious phase of the life cycle of the plant, as in the
mosses, or it can occur in a microscopic structure, such as a pollen
grain, in the higher plants (a common collective term for the
vascular plants).
• The sporophyte stage is barely noticeable in lower plants (the
collective term for the plant groups of mosses, liverworts, and
lichens). Towering trees are the diplontic phase in the lifecycles of
plants such as sequoias and pines.
• Protection of the embryo is a major requirement for land plants.
• The vulnerable embryo must be sheltered from desiccation and
other environmental hazards. In both seedless and seed plants, the
female gametophyte provides protection and nutrients to the
embryo as it develops into the new generation of sporophyte. This
distinguishing feature of land plants gave the group its alternate
name of embryophytes.
 Sporangia in Seedless Plants
• The sporophyte of seedless plants is diploid
and results from syngamy (fusion) of two
gametes.
• The sporophyte bears the sporangia
(singular, sporangium): organs that first
appeared in the land plants.
• The term “sporangia” literally means “spore
in a vessel,” as it is a reproductive sac that
contains spores
• Two different types of spores are produced in
land plants, resulting in the separation of
sexes at different points in the
lifecycle. Seedless non-vascular
plants produce only one kind of spore and
are called homosporous. The
gametophyte phase is dominant in these
plants. After germinating from a spore, the
resulting gametophyte produces both male
and female gametangia, usually on the same
individual
• In contrast, heterosporous plants produce two morphologically
different types of spores. The male spores are called microspores,
because of their smaller size, and develop into the male
gametophyte; the comparatively larger megaspores develop into
the female gametophyte. Heterospory is observed in a few seedless
vascular plants and in all seed plants.
• When the haploid spore germinates in a hospitable environment, it
generates a multicellular gametophyte by mitosis.
• The spores of seedless plants are surrounded by thick cell walls
containing a tough polymer known as sporopollenin.
• Sporopollenin is unusually resistant to chemical and biological
degradation. In seed plants, which use pollen to transfer the male
sperm to the female egg, the toughness of sporopollenin explains the
existence of well-preserved pollen fossils.
• Sporopollenin was once thought to be an innovation of land plants;
however, the green algae Coleochaetes forms spores that contain
sporopollenin.
Gametangia in Seedless Plants

• Gametangia (singular, gametangium) are structures observed on

multicellular haploid gametophytes.
• The male gametangium (antheridium) releases sperm. Many
seedless plants produce sperm equipped with flagella that enable
them to swim in a moist environment to the archegonia: the
female gametangium.
• Gametangia are prominent in seedless plants, but are very rarely
found in seed plants.
Apical Meristems
Shoots and roots of plants increase in
length through rapid cell division in a
tissue called the apical meristem, which
is a small zone of cells found at the shoot
tip or root tip. The apical meristem is
made of undifferentiated cells that
continue to proliferate throughout the
life of the plant. Meristematic cells give
rise to all the specialized tissues of the
organism. Elongation of the shoots and
roots allows a plant to access additional
space and resources: light in the case of
the shoot, and water and minerals in the
case of roots. A separate meristem, called
the lateral meristem, produces cells that
increase the diameter of tree trunks.
Additional Land Plant Adaptations
• As plants adapted to dry land and became independent from the
constant presence of water in damp habitats, new organs and
structures made their appearance.
• Early land plants did not grow more than a few inches off the
ground, competing for light on these low mats. By developing a
shoot and growing taller, individual plants captured more light.
• In small plants such as single-celled algae, simple diffusion suffices
to distribute water and nutrients throughout the organism.
However, for plants to evolve larger forms, the evolution of vascular
tissue for the distribution of water and solutes was a prerequisite.
• The vascular system contains xylem and phloem tissues. Xylem
conducts water and minerals absorbed from the soil up to the shoot,
while phloem transports food derived from photosynthesis
throughout the entire plant.
• In land plants, a waxy, waterproof cover called a cuticle protects the
leaves and stems from desiccation. However, the cuticle also
prevents intake of carbon dioxide needed for the synthesis of
carbohydrates through photosynthesis. To overcome this, stomata
or pores that open and close to regulate traffic of gases and water
vapor appeared in plants as they moved away from moist
environments into drier habitats.
• Water filters ultraviolet-B (UVB) light, which is harmful to all
organisms, especially those that must absorb light to survive. This
filtering does not occur for land plants. This presented an additional
challenge to land colonization, which was met by the evolution of
biosynthetic pathways for the synthesis of protective flavonoids and
other compounds: pigments that absorb UV wavelengths of light
and protect the aerial parts of plants from photodynamic damage.
• Plants cannot avoid being eaten by animals. Instead, they synthesize
a large range of poisonous secondary metabolites: complex organic
molecules such as alkaloids, whose noxious smells and unpleasant
taste deter animals.
Evolution of Land Plants • The early era, known as the Paleozoic,
is divided into six periods. It starts
with the Cambrian period, followed
by the Ordovician, Silurian, Devonian,
Carboniferous, and Permian.
• The major event to mark the
Ordovician, more than 500 million
years ago, was the colonization of
land by the ancestors of modern land
plants. Fossilized cells, cuticles, and
spores of early land plants have been
dated as far back as the Ordovician
period in the early Paleozoic era.

• The oldest-known vascular plants have been identified in deposits from

the Devonian. One of the richest sources of information is the Rhynie
chert, a sedimentary rock deposit found in Rhynie, Scotland (Figure 5),
where embedded fossils of some of the earliest vascular plants have been
identified.
PALEOBOTANIST
• Paleobotanists distinguish between extinct species, as fossils,
and extant species, (the study of extinct plants).
• Paleobotanists trace the evolution of plants by following the modifications
in plant morphology: shedding light on the connection between existing
plants by identifying common ancestors that display the same traits.
• Paleobotanists collect fossil specimens in the field and place them in the
context of the geological sediments and other fossilized organisms
surrounding them. The activity requires great care to preserve the integrity
of the delicate fossils and the layers of rock in which they are found.
• Use of analytical chemistry and molecular biology to study fossils.
• One example of the use of analytical chemistry and molecular biology is the
identification of oleanane, a compound that deters pests. Up to this point,
oleanane appeared to be unique to flowering plants; however, it has now
been recovered from sediments dating from the Permian, much earlier than
the current dates given for the appearance of the first flowering plants.
• Paleobotanists can also study fossil DNA, which can yield a large
amount of information, by analyzing and comparing the DNA
sequences of extinct plants with those of living and related
organisms.
• Some paleobotanists are skeptical of the conclusions drawn from
the analysis of molecular fossils
• For example, the chemical materials of interest degrade rapidly
when exposed to air during their initial isolation, as well as in
further manipulations.
The Major Divisions of Land Plants
• The green algae and land plants are grouped together into a
subphylum called the Streptophytina, and thus are called
Streptophytes.
• In a further division, land plants are classified into two major
groups according to the absence or presence of vascular tissue.
• Plants that lack vascular tissue, which is formed of specialized cells
for the transport of water and nutrients, are referred to as non-
vascular plants. Liverworts, mosses, and hornworts are seedless,
non-vascular plants that likely appeared early in land plant
evolution.
• Vascular plants developed a network of cells that conduct water and
solutes. The first vascular plants appeared in the late Ordovician
and were probably similar to lycophytes, which include club mosses
(not to be confused with the mosses) and the pterophytes (ferns,
horsetails, and whisk ferns).
• Lycophytes and pterophytes are referred to as seedless vascular
plants, because they do not produce seeds.
• The seed plants, or spermatophytes, form the largest group of all
existing plants, and hence dominate the landscape. Seed plants
include gymnosperms, most notably conifers (Gymnosperms),
which produce “naked seeds,” and the most successful of all plants,
the flowering plants (Angiosperms). Angiosperms protect their
seeds inside chambers at the center of a flower; the walls of the
chamber later develop into a fruit.
Green Algae: Precursors of Land Plants
Streptophytes
• Until recently, all photosynthetic eukaryotes were considered
members of the kingdom Plantae.
• The brown, red, and gold algae, however, have been reassigned to
the Protista kingdom. This is because apart from their ability to
capture light energy and fix CO, they lack many structural and
biochemical traits that distinguish plants from protists.
• The position of green algae is more ambiguous. Green algae contain
the same carotenoids and chlorophyll a and b as land plants,
whereas other algae have different accessory pigments and types of
chlorophyll molecules in addition to chlorophyll a. Both green algae
and land plants also store carbohydrates as starch. Cells in green
algae divide along cell plates called phragmoplasts, and their cell
walls are layered in the same manner as the cell walls of
embryophytes. Consequently, land plants and closely related green
algae are now part of a new monophyletic group called
Streptophyta.
• The remaining green algae, which belong to a group called Chlorophyta,
include more than 7000 different species that live in fresh or brackish
water, in seawater, or in snow patches.
• A few green algae even survive on soil, provided it is covered by a thin film
of moisture in which they can live.
• Periodic dry spells provide a selective advantage to algae that can survive
water stress. Some green algae may already be familiar, in
particular Spirogyra and desmids. Their cells contain chloroplasts that
display a dizzying variety of shapes, and their cell walls contain cellulose, as
do land plants.
• Some green algae are single cells, such as Chlorella and Chlamydomonas,
which adds to the ambiguity of green algae classification, because plants are
multicellular. Other algae, like Ulva (commonly called sea lettuce), form
colonies
Reproduction of Green Algae

• Green algae reproduce both asexually, by fragmentation or dispersal

of spores, or sexually, by producing gametes that fuse during
fertilization. In a single-celled organism such as Chlamydomonas,
there is no mitosis after fertilization. In the multicellular Ulva, a
sporophyte grows by mitosis after fertilization.
Both Chlamydomonas and Ulva produce flagellated gametes.
Charales • Green algae in the order Charales,
and the coleochaetes (microscopic
green algae that enclose their
spores in sporopollenin), are
considered the closest living
relatives of embryophytes.
• The Charales can be traced back
420 million years. They live in a
range of fresh water habitats and
vary in size from a few
millimeters to a meter in length.
• The representative species
is Chara, often called muskgrass
or skunkweed because of its
unpleasant smell.
• Unlike land plants, Charales do
not undergo alternation of
generations in their lifecycle.
• Charales exhibit a number of traits that are significant in their
adaptation to land life. They produce the compounds lignin and
sporopollenin, and form plasmodesmata that connect the cytoplasm
of adjacent cells. The egg, and later, the zygote, form in a protected
chamber on the parent plant.
Bryophytes
• group of plants that are the closest extant relative of early
terrestrial plants.
• The first bryophytes (liverworts) most likely appeared in the
Ordovician period, about 450 million years ago.
• More than 25,000 species of bryophytes thrive in mostly damp
habitats, although some live in deserts.
• Although the term non-tracheophyte is more accurate,
bryophytes are commonly called nonvascular plants.
• In a bryophyte, all the conspicuous vegetative organs—including
the photosynthetic leaf-like structures,the thallus, stem, and the
rhizoid that anchors the plant to its substrate—belong to the
haploid organism or gametophyte. The sporophyte is barely
noticeable.
• The multicellular sexual reproductive structure is present in
bryophytes and absent in the majority of algae.
• The bryophytes are divided into three phyla: the liverworts or
Hepaticophyta, the hornworts or Anthocerotophyta, and the
mosses or true Bryophyta.
Liverworts
(Hepaticophyta)
• are viewed as the
plants most closely
related to the ancestor
that moved to land.
• Liverworts have
colonized every
terrestrial habitat on
Earth and diversified
to more than 7000
existing species.
• Openings that allow
the movement of
gases may be observed
in liverworts
• Represents the lifecycle of a liverwort. The cycle
starts with the release of haploid spores from the
sporangium that developed on the sporophyte.
Hornworts
(Anthocerotophyta)
• Hornworts belong to the broad
bryophyte group. They have
colonized a variety of habitats on
land, although they are never far
from a source of moisture.
• The short, blue-green
gametophyte is the dominant
phase of the lifecycle of a
hornwort.
• The narrow, pipe-like sporophyte
is the defining characteristic of the
group.
• Many hornworts establish
symbiotic relationships with
cyanobacteria that fix nitrogen
from the environment
• The lifecycle of hornworts follows the general pattern
of alternation of generations.
Mosses
• More than 10,000 species of mosses have been catalogued.
Their habitats vary from the tundra, wherethey are the main
vegetation, to the understory of tropical forests. In the
tundra, the mosses’ shallow rhizoids allow them to fasten to a
substrate without penetrating the frozen soil.
• Mosses slow down erosion, store moisture and soil nutrients,
and provide shelter for small animals as well as food for
larger herbivores, such as the musk ox.
• Mosses are very sensitive to air pollution and are used to
monitor air quality. They are also sensitive to copper salts, so
these salts are a common ingredient of compounds marketed
to eliminate mosses from lawns.
• Mosses form diminutive gametophytes, which are the
dominant phase of the lifecycle. Green, flat Structures
resembling true leaves, but lacking vascular tissue—are
attached in a spiral to a central stalk.
The moss lifecycle follows the pattern of alternation of generations as
shown in Figure 25.14. The most familiar structure is the haploid
gametophyte, which germinates from a haploid spore and forms first a
protonema usually,a tangle of single-celled filaments that hug the
ground. Cells akin to an apical meristem actively divide and give rise to
a gametophore, consisting of a photosynthetic stem and foliage-like
structures
Seedless Vascular Plants
• The vascular plants, or tracheophytes, are the
dominant and most conspicuous group of land plants.
• More than 260,000 species of tracheophytes
represent more than 90 percent of Earth’s vegetation.
• Several evolutionary innovations explain their success
and their ability to spread to all habitats.
• Vascular plants, on the other hand, can achieve
enormous heights, thus competing successfully for
light. Photosynthetic organs become leaves, and pipe-
like cells or vascular tissues transport water, minerals,
and fixed carbon throughout the organism
Vascular Tissue: Xylem and Phloem
• The first fossils that show the presence of vascular tissue date to the Silurian
period, about 430 millionyears ago. The simplest arrangement of conductive
cells shows a pattern of xylem at the center surrounded by phloem.
• Xylem is the tissue responsible for the storage and long-distance transport
ofwater and nutrients, as well as the transfer of water-soluble growth factors
from the organs of synthesis to the target organs.
• The tissue consists of conducting cells, known as tracheids, and supportive
filler tissue, called parenchyma.
• Xylem conductive cells incorporate the compound lignin into their walls, and
are thus described as lignified. Lignin itself is a complex polymer that is
impermeable to water and confers mechanical strength to vascular tissue.
• With their rigid cell walls, the xylem cells provide support to the plant and
allow it to achieve impressive heights. Tall plants have a selective advantage by
being able to reach unfiltered sunlight and disperse their spores or seeds
further away, thus expanding their range.
• Phloem is the second type of vascular tissue; it transports sugars, proteins, and
other solutes throughout the plant.
• Phloem cells are divided into sieve elements (conducting cells) and cells that
support the sieve elements. Together, xylem and phloem tissues form the
vascular system of plants.
Roots: Support for the Plant
• Roots are not well preserved in the fossil record.
Nevertheless, it seems that roots appeared laterv in
evolution than vascular tissue.
• Thin rhizoids attached bryophytes to the substrate,
but these rather flimsy filaments did not provide a
strong anchor for the plant; neither did they absorb
substantial amounts of water and nutrients.
• In contrast, roots, with their prominent vascular
tissue system, transfer water and minerals from the
soil to the rest of the plant.
• The majority of roots establish a symbiotic
relationship with fungi, forming mycorrhizae, which
benefit the plant by greatly increasing the surface
area for absorption of water and soil minerals and
nutrients.
Leaves, Sporophylls, and Strobili
• A third innovation marks the seedless vascular plants. Accompanying the
prominence of the sporophyte and the development of vascular tissue, the
appearance of true leaves improved their photosynthetic efficiency.
• Leaves capture more sunlight with their increased surface area by employing
more chloroplasts to trap light energy and convert it to chemical energy, which
is then used to fix atmospheric carbon dioxide into carbohydrates. The
carbohydrates are exported to the rest of the plant by the conductive cells of
phloem tissue.
• The first type of leaf is the microphyll, or “little leaf,” which can be dated to 350
million years ago in the late Silurian.
• A microphyll is small and has a simple vascular system. A single unbranched
vein—a bundle of vascular tissue made of xylem and phloem—runs through the
center of the leaf.
• Megaphylls most likely appeared independently several times during the course
of evolution. Their complex networks of veins suggest that several branches may
have combined into a flattened organ, with the gaps between the branches being
filled with photosynthetic tissue.
• Strobili are cone-like structures that contain sporangia. They are prominent in
conifers and are commonly known as pine cones.
Ferns and Other Seedless Vascular
Plants
• By the late Devonian period, plants had evolved vascular
tissue, well-defined leaves, and root systems.
• With these advantages, plants increased in height and
size. During the Carboniferous period, swamp forests of
club mosses and horsetails some specimens reaching
heights of more than 30 m (100ft) covered most of the
land.
• In seedless vascular plants, the sporophyte became the
dominant phase of the lifecycle.
• Water is still required for fertilization of seedless
vascular plants, and most favor a moist environment.
• Modern-day seedless tracheophytes include club
mosses, horsetails, ferns, and whisk ferns.
Phylum Lycopodiophyta:
Club Mosses
• The club mosses, or phylum
Lycopodiophyta, are the earliest
group of seedless vascular
plants. Theydominated the
landscape of the Carboniferous,
growing into tall trees and
forming large swamp forests.
• Lycophytes follow the pattern of
alternation of generations seen
in the bryophytes, except that
the sporophyte is the major
stage of the lifecycle.
• Lycophytes can be homosporous
or heterosporous.
 Phylum Monilophyta: Class
Equisetopsida (Horsetails)
• Horsetails, whisk ferns and ferns belong to
the phylum Monilophyta, with horsetails
placed in the Class Equisetopsida.
• The single genus Equisetum is the survivor
of a large group of plants, known as
Arthrophyta, which produced large trees
and entire swamp forests in the
Carboniferous. The plants are usually
found in damp environments and marshes.
• Silica collects in the epidermal cells,
contributing to the stiffness of horsetail
plants. Underground stems known as
rhizomes anchor the plants to the ground.
Modern-day horsetails are homosporous
and produce bisexual gametophytes.
Phylum Monilophyta:
Class Psilotopsida
(Whisk Ferns)
• While most ferns form large leaves
and branching roots, the whisk
ferns, Class Psilotopsida, lack both
roots and leaves, probably lost by
reduction.
• Photosynthesis takes place in their
green stems, and small yellow
knobs form at the tip of the branch
stem and contain the sporangia.
• Whisk ferns were considered an
early pterophytes.
Phylum Monilophyta: Class
Psilotopsida (Ferns)
• With their large fronds, ferns are the
most readily recognizable seedless
vascular plants. They are considered the
most advanced seedless vascular plants
and display characteristics commonly
observed in seed plants.
• Ferns made their appearance in the fossil
record during the Devonian period and
expanded during the Carboniferous.
• The dominant stage of the lifecycle of a
fern is the sporophyte, which consists of
large compound leaves called fronds.
Fronds fulfill a double role; they are
photosynthetic organs that also carry
reproductive organs.
• The tip of a developing fern frond is
rolled into a crozier, or fiddlehead.

• Fiddleheads unroll as the frond develops.

Life Cycle of Fern

Life Cycle of Fern

Most ferns produce the same type of spores and
are therefore homosporous.
The diploid sporophyte is the most conspicuous
stage of the lifecycle. On the underside of its
mature fronds, sori (singular, sorus)
form as small clusters where sporangia develop.

• Inside the sori, spores are produced by

meiosis and released into the air. Those
that land on a suitable substrate
germinate and form a heart-shaped
gametophyte, which is attached to the
ground by thin filamentous rhizoids.
• The inconspicuous gametophyte harbors
both sex gametangia. Flagellated sperm
released from the antheridium swim on a
wet surface to the archegonium, where the
egg is fertilized.
• The newly formed zygote grows into a
sporophyte that emerges from the
gametophyte and grows by mitosis into the
next generation sporophyte.
The Importance of Seedless Vascular Plants
• Mosses and liverworts are often the first macroscopic organisms to colonize
an area, both in a primary succession where bare land is settled for the first
time by living organisms or in a secondary succession, where soil remains
intact after a catastrophic event wipes out many existing species.
• Their spores are carried by the wind, birds, or insects. Once mosses and
liverworts are established, they provide food and shelter for other species. In
a hostile environment, like the tundra where the soil is frozen, bryophytes
grow well because they do not have roots and can dry and rehydrate rapidly
once water is again available.
• Mosses are at the base of the food chain in the tundra biome. Many species
fromsmall insects to musk oxen and reindeer depend on mosses for food. In
turn, predators feed on the herbivores, which are the primary consumers.
• At the end of the nineteenth century, scientists observed that lichens and
mosses were becoming increasingly rare in urban and suburban areas. Since
bryophytes have neither a root system for absorption of water and nutrients,
nor a cuticle layer that protects them from desiccation, pollutants in
rainwater readily penetrate their tissues; they absorb moisture and nutrients
through their entire exposed surfaces.
• Therefore, pollutants dissolved in rainwater penetrate plant tissues
readily and have a larger impact on mosses than on other plants. The
disappearance of mosses can be considered a bioindicator for the level of
pollution in the environment.
• Ferns contribute to the environment by promoting the weathering of
rock, accelerating the formation of topsoil, and slowing down erosion by
spreading rhizomes in the soil. The water ferns of the genus Azolla harbor
nitrogen-fixing cyanobacteria and restore this important nutrient to
aquatic habitats.
• Seedless plants have historically played a role in human life through uses
as tools, fuel, and medicine.Dried peat moss, Sphagnum, is commonly
used as fuel in some parts of Europe and is considered a renewable
resource. Sphagnum bogs (Figure 25.26) are cultivated with cranberry
and blueberry bushes.
The ability of Sphagnum to hold moisture makes
the moss a common soil conditioner. Florists
use blocks of Sphagnum to maintain moisture
for floral arrangements.
Americans in the Pacific Northwest, and are popular as a
side dish in French cuisine. The licorice fern,

Polypodium glycyrrhiza, is part of the diet of the Pacific

Northwest coastal tribes, owing in part to the sweetness of
its rhizomes. It has a faint licorice taste and serves as a
sweetener. The rhizome also figures in the pharmacopeia
of Native Americans for its medicinal properties and is
used as a remedy for sore throat.