Copyright © Fall 2024, Biology 1B, University of California Berkeley

Lab 8 – Terrestrialization
What to do before lab:

● Read this lab introduction BEFORE coming to lab and familiarize yourself with the concepts
● Complete the Pre-Lab Videos and Assignments
● Review and define the list of important terminology (see end of this document)
● Review the Organismal Biology lectures
● Review mitosis and meiosis in your text. Focus on the ploidy of the initial and the
resulting cells in each process. Details about the specific phases of each process are not necessary
for the Biology 1B labs.

Things to bring to lab:

● Bring this introduction and your list of important terms (above) OR digital access to these.
● Bring your textbook or digital access to the e-text (see bCourses for access instructions)

Cyanobacteria
Overview
Cyanobacteria are microscopic organisms that are common in aquatic and terrestrial ecosystems. They are named for the
bluish pigment phycocyanin which they use to capture light energy in photosynthesis (although not all cyanobacteria are
blue). Cyanobacteria are abundant components of marine and freshwater phytoplankton, the collection of
photosynthetic organisms that float near the surface of a body of water. Phytoplankton is ecologically essential,
accounting for much of the primary production (carbon capture as organic molecules) that occurs on Earth.

Figure 1 (left). Cyanobacteria cells may be organized into filaments.

Figure 2 (right) . Diagram of a cyanobacterium. Image shows one unicellular cyanobacterial organism that is part of a filament of
multiple cyanobacterial cells. Note the presence of cellular structures, but absence of membrane-bound organelles or nucleus.

Prokaryotes vs. Eukaryotes

Cyanobacteria are the only prokaryotes that you will observe in Bio 1B labs. Therefore, it is useful to contrast
prokaryotes and eukaryotes (all of the other organisms that you will observe in Bio 1B labs are eukaryotes). Prokaryotes
are single-celled organisms in the domains Archaea and Bacteria. While there are both unicellular and multicellular
eukaryotes, prokaryotes are only unicellular. Prokaryotic cells are generally smaller than eukaryotic cells and the
structure of prokaryotic cells differs in important ways. One of the most important differences is that prokaryotes lack
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
organelles, specialized membrane-bound structures inside of the cell such as nuclei, mitochondria, and chloroplasts.
Moreover, the structure of prokaryotic cell walls differs from the structure of eukaryotic cell walls (in those eukaryotes
that have cell walls). Finally, prokaryotes do not reproduce sexually; they reproduce asexually through binary fission (a
form of cell division).

Cyanobacteria are photoautotrophs

Autotrophs are organisms that produce their own carbon-based food. Heterotrophs are organisms that acquire their
carbon-based food by consuming other organisms or substances produced by other organisms (heterotrophs include
animals, fungi, etc.). Cyanobacteria are photosynthetic autotrophs (or ‘photoautotrophs’): they use carbon dioxide and
light energy from the sun to produce their own carbon-based food (glucose). Cyanobacteria are the only
oxygen-producing photosynthetic prokaryotes. Oxygen is a by-product of their photosynthesis, and cyanobacteria have
important ecological roles as oxygen producers. Cyanobacteria are thought to have dramatically increased oxygen levels
in the Earth’s atmosphere causing the Great Oxygenation Event more than two billion years ago.

Endosymbiosis
Although cyanobacteria have photosynthetic pigments, the pigments are not contained within organelles—they have no
organelles. Chloroplasts are the organelles in green plants (eukaryotes) that contain chlorophyll, the light-capturing
pigment necessary for photosynthesis in green plants. Chloroplasts likely originated as symbiotic cyanobacteria that were
incorporated into the cells of ancient eukaryotes. This process of incorporation of one lineage into the cells of another
lineage is called endosymbiosis. These cyanobacteria became organelles in the cells of these ancient eukaryotes.

Cellular organization
Like all prokaryotes, cyanobacteria are single-celled organisms. Some cyanobacteria exist in a purely unicellular form,
while others aggregate into colonies. Colonies can take the form of filaments, sheets, or hollow spheres. These colonial
forms allow some differentiation of function among individual cells that make up the colony, but they do not represent
true multicellularity. For example, the filamentous colonies of some types of cyanobacteria contain cells called
heterocysts which are specialized for nitrogen fixation, the process whereby the nitrogen from nitrogen gas is ‘fixed’ into
a solid form. Some cyanobacteria live in symbiotic relationships with fungi (lichen) and with plants, and they become
incorporated into the
structure of these
organisms.
Figure 3 & 4. Examples of
filamentous cyanobacterial
colonies.

Figure 5 (right). Dihu

(Spirulina) harvesting and
processing in Chad.

Dietary use
Spirulina is a type of
cyanobacterium that is sold
as a dietary supplement. Spirulina is high in protein and has been an important food source in some indigenous cultures.
For example, Spirulina (called tecuitlatl) harvested from Lake Texcoco (site of present-day Mexico City) was an important
food source for the Mexicas who inhabited this area. Spirulina (called dihu) is also harvested by the Kanembu people
from Lake Chad in West-central Africa.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley

Cyanobacterial blooms
Fluctuation in environmental factors such as light availability, water temperature, water flow, pH, and nitrogen and
phosphorous levels can lead to fluctuations in the sizes of cyanobacteria populations. If cyanobacteria populations get
too large, they can create cyanobacterial blooms, which can cause serious problems for aquatic ecosystems. Numerous
types of cyanobacteria produce cyanotoxins, toxic substances which can be harmful (or even deadly) to wildlife,
livestock, and humans.

Eutrophication, a process in which nutrients, particularly phosphorus and nitrogen, become highly concentrated in a
body of water, can be a key driver in cyanobacterial blooms. Eutrophication can have harmful effects on the organisms
that inhabit aquatic ecosystems because it reduces the amount of dissolved oxygen in the water, which can lead to the
death of aquatic plants and animals. Eutrophication involving cyanobacteria can be dangerous because cyanobacteria
can produce a range of potent toxins, making some blooms deadly to animal life (including humans).

The frequency of cyanobacterial blooms has been increasing due to anthropogenic activities such as discharges from
wastewater treatment plants or municipal sewer systems, raising of livestock, and runoff from agricultural fields and
roads, which can be high in nitrogen and phosphorus.

Figure 6. Anthropogenic contributions to cyanobacterial blooms

Figure 7 (left). Eutrophication at a waste water outlet in the

Potomac River, Washington D.C. (Photo: Trubetskoy)Figure 8 (right).
Cyanobacterial bloom in the Pacific Ocean near Fiji, as seen from
space (NASA).

Volvox
Overview
Volvox are colonial freshwater green algae. They are
eukaryotes, and belong to a group that includes red algae,
green algae, and land plants. Volvox are common in
freshwater ecosystems throughout the world. Volvox are
photosynthetic, and it is the green photosynthetic
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
chlorophyll pigment in their chloroplasts that gives them their green color. Volvox colonies serve as an important food
source for aquatic microorganisms such as paramecia and rotifers.
Figure 9. Volvox colonies with daughter colonies visible inside.

Cellular organization
Volvox colonies are hollow spheres of cells embedded in an extracellular gelatinous protein matrix. The cells are usually
connected within the colony by cytoplasmic bridges. These bridges are composed of cytoplasm that is shared between
neighboring cells and are the result of incomplete cell division. The bridges allow cells to share certain organelles, like
mitochondria, and also allow a form of communication between different cells of the colony. This aids in the coordinated
movement of the individual unicellular organisms that make up the colony and allows them to coordinate reproduction.
This kind of coordination between different cells is similar to what we find in truly multicellular organisms.

The movement of Volvox colonies is the result of coordinating the flagella of each individual cell on the colony’s surface.
These colonies do not have a nervous system, and exactly how they coordinate flagella movement is not
well-understood.

Volvox colonies contain two distinct cell types

Volvox colonies contain two distinct cell types: somatic cells and germ cells. Somatic cells (non-reproductive cells of the
body) have two flagella pointed outward and help to move the colony in a coordinated fashion toward light. Because
they are photosynthetic, Volvox need to be exposed to sunlight. Somatic cells have eyespots (visible as a small red dot in
the cell) that allows phototaxis, the movement of the organism in response to light. Volvox colonies have distinct anterior
and posterior poles and the somatic cells closer to the anterior pole have more developed eyespots. This helps the
colony to swim toward the light. Somatic cells will eventually die without dividing. There is also a much smaller number
of germ cells which aid in reproduction.

Figure 10 (left). Diagram of the structure of a Volvox colony. 1. Single Volvox cell. 2. Daughter colony. 3. Cytoplasmic bridges. 4.
Intercellular gelatinous matrix. 5. Reproductive cell. 6. Somatic cell.

Figure 11 (right). Closeup of surface of Volvox colony. Note the individual unicellular organisms that make up the colony. Cytoplasmic
bridges between cells are visible near the top of the colony.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
Figure 12 (left). Close-up of the individual cells of a Volvox colony. The two flagella and the red eyespot are visible for some of the
individual cells.

Figure 13 (right). Close-up of the surface of a Volvox colony with a large daughter colony in the center. Note the thin cytoplasmic
bridges that connect the individual cells.

Volvox can reproduce asexually or sexually

Asexual reproduction results in the production of daughter colonies by larger colonies through the process of mitosis.
These colonies mature within the parent colony, then burst out of the parent colony when they are fully mobile,
destroying the parent colony in the process.

The change from asexual to sexual reproduction occurs due to environmental stress-–for example, if the environment
starts to dry out. Sexual reproduction increases genetic diversity and is advantageous when environmental conditions are
less stable. In sexual reproduction, sperm fertilize eggs, resulting in the production of zygotes that may resist
environmental stress. The zygotes can then germinate (begin to grow into new colonies) when environmental conditions
become more suitable.

Figure 14. Left: Volvox with daughter colonies

within a parent colony (photo: Hallmann).
Right: Daughter colonies being released from a
parent colony.

Multicellular green algae

Many botanical terms come from Greek or Latin, and they are created by adding prefixes or suffixes. For example, -phyte
means plant; by adding gameto-, it becomes gametophyte, a plant that produces gametes. Using the same logic, a
sporophyte is a plant that produces spores. Knowing word origins is useful when learning scientific terms. Learning the
list below will be a tremendous aid in learning the material in the organismal diversity labs.

Tips for learning botanical terms:

“homo-” – equal, or same (homosporous) “-phyll” – leaf (sporophyll)

“hetero-” – different (heterosporous) “-angium” – vessel or container (sporangium,

gametangium)
“-phyte” – plant (sporophyte, gametophyte)
“-sperm” – seed (gymnosperm, angiosperm)
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
“mega-” – big (megagametophyte, megaspore) “mei-" – to lessen (meiosis)

“micro-” – small (microgametophyte, microspores) “anth-” – male (anther, antheridia)

“-cyte” – cell (sporocyte) “arch-” – female (archegonium)

Ulva
Ulva (also called ‘sea lettuce’) is a type of multicellular green
alga that grows in coastal aquatic environments. It is
eukaryotic, photosynthetic, and truly multicellular like other
plants. The body is composed of two layers of cells that form
the thallus, the leaf-like body of the individual. The lower
part of the individual develops into a structure called a
holdfast that anchors the plant to the substrate. Ulva differs
from Volvox in that it is sessile as an adult (it cannot move or
change position). It is a truly multicellular organism because
it has permanently associated tissues (i.e., tissues that do not
exist separately) with different structures and functions that
communicate to coordinate their activities.

Ulva has a haplodiplontic life cycle (also called “alternation of generations”), a characteristic also found in land plants.
The life cycles are named for the multicellular generations that are present. In the haplodiplontic life cycle, there are
two multicellular generations: a multicellular diploid sporophyte and a multicellular haploid gametophyte. The
sporophyte and the gametophyte are two different multicellular organisms. In Ulva, the sporophyte and the
gametophyte look identical. In other words, the two generations are isomorphic, meaning they have the “same
morphology.”

The sporophytes produce haploid spores via meiosis. In Ulva, the spores are able to swim by means of flagella. These
spores grow by mitosis into gametophytes. The mature gametophytes produce haploid gametes via mitosis. Both types
of gamete are released directly into the water around the plant (i.e., they are not protected in structures on the
gametophyte, as we will see in some other plants). The fusion of gametes produces a single-celled, diploid zygote. The
zygote will develop into a new multicellular sporophyte through mitosis. The cycle will begin again as these sporophytes
produce spores.

All land plants have the haplodiplontic life cycle. You probably learned in human biology that we (like all animals)
produce gametes (sperm and egg) via meiosis. It is important to remember that plants produce gametes via mitosis.
Plants produce spores via meiosis.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
Figure 16. The Ulva life cycle, typical of all haplodiplontic life cycles. The generations alternate between diploid sporophytes and
haploid gametophytes. Although two types of gametophytes are produced in
the Ulva life cycle, Ulva is not heterosporous
(they produce only one type of spore).

Chara
Charophytes (including the genus Chara)
are multicellular green algae that live in
freshwater. We will use the genus Chara
to highlight some of the characteristics of
freshwater green algae. Charophytes are
thought to be the closest relatives of
land plants. Charophytes share
distinctive traits with land plants that suggest this close relationship. For example, the sperm flagella of charophytes and
land plants is similar (the same flagella structure is not found in other groups of green algae). In addition, both
charophytes and land plants retain the zygotes on the parent plant.

Figure 17. Chara is a multicellular plant that lives in fresh water. It has specialized structures to protect gametes.

Charophyte gametophytes have specialized structures that protect the two types of gametes: eggs and sperm. The sperm
is enclosed in a structure called an antheridium (plural, antheridia). The egg is produced in a structure called an
oogonium (plural, oogonia). The sperm are eventually released from the antheridium, but the egg is never released.
Sperm released from the antheridium swim through water to fertilize the egg in the oogonium. (Although the protective
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
function of the oogonium is similar to the protective function of the archegonium that you will observe later in land
plants, they are different structures).

Oogonia can be identified by their location at the nodes (the points where the leaf-like branchlets meet the stem), by
their spiral outer structure, and by the crown-shaped opening at the top end. Antheridia are also located at the nodes,
but they are sphere-shaped rather than spiral-shaped.

Chara have a haplontic life cycle. Remember: the life cycles are named for the multicellular generations that are present.
In the haplontic life cycle, there is only one multicellular generation: the haploid gametophyte. There is a diploid phase
in the life cycle (the zygote), but it is not multicellular. As in all of the plants we study, the single-celled zygote is produced
by the fertilization of the egg by the sperm. Unlike organisms with a haplodiplontic life cycle, however, the zygote does
not grow via mitosis to become a multicellular sporophyte. Rather, the single-celled zygote immediately undergoes
meiosis to produce haploid spores.

Figure 19. Diagram of haplontic life cycle of charophyte.

Land plant life cycle

Overview
There are three major groups of land plants: bryophytes (e.g., mosses), pteridophytes (e.g., ferns) and seed plants
(e.g., conifers and flowering plants). The diversity of land plants is incredible, ranging from tiny mosses that grow on
mountain tops to giant trees in tropical forests, from beautiful water lilies to desert cacti. Despite all of this diversity, all
land plants have a haplodiplontic life cycle.

The haplodiplontic life cycle (alternation of generations)

The haplodiplontic life cycle is characterized by an alternation of two different multicellular generations: the
sporophyte generation and the gametophyte generation. (When we use the term “generation,” we are always referring
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
to a multicellular plant). Sporophytes are multicellular diploid plants. Gametophytes are multicellular haploid plants.
Thus, all land plants have both gametophytes and sporophytes, but the two generations are not always easy to see. For
example, when you are looking at a redwood forest you are looking at giant redwood sporophytes, and when you are
looking at a green moss carpeting a rock, you are looking at moss gametophytes. Where are the gametophytes of the
redwoods and the sporophytes of the mosses?

Figure 20. The carpet-like moss is a gametophyte, while the giant redwood tree is a sporophyte.

The following image is a representation of the haplodiplontic life cycle.

Figure 21. The haplodiplontic life cycle.

Variations of the haplodiplontic life cycle

Although the basic characteristics of the life cycle of all land plants are essentially the same, there is variation among the
three major groups in the following characteristics:

Dominance. One of the multicellular generations (gametophyte or sporophyte) is larger (has more biomass), can live for
a longer amount of time, and can generate offspring over multiple seasons. We refer to this as the dominant generation.
For example, in a life cycle in which the sporophyte is dominant, the sporophyte may live for several years and produce
offspring (gametophytes) multiple times, whereas the gametophytes (which are not dominant) may live only for a few
days and produce only one generation of sporophyte offspring. The dominant generation is also larger and more
conspicuous (clearly visible) and generally has a more complex morphology.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
Nutritional dependence. Either the gametophyte or the sporophyte may lack the ability to perform photosynthesis.
Instead, it acquires its carbon-based food from its parent (which can be either the sporophyte or the gametophyte,
depending on the taxonomic group). If a generation can produce all of its own carbon-based food through
photosynthesis, we say that the generation is independent. If it acquires much or all of its carbon-based food from the
other generation, we say that it is dependent. For example, in bryophytes, the gametophyte is independent, while the
sporophyte is dependent on the gametophyte. In flowering plants, the gametophyte is dependent on the sporophyte,
while the sporophyte is independent. If a generation is dependent, it will lack green pigments (chlorophyll) and have a
physical connection to the other
generation (e.g., the bryophyte
sporophyte grows out of, and is
connected to, the gametophyte).

Figure 22. Left: Moss life cycle. The gametophyte is

dominant. The sporophyte is dependent on the
gametophyte. Right: Fern life cycle. The
sporophyte is dominant, but both generations
are independent.

Dependence on water for fertilization. In some land plants, the male gametes (sperm) have to swim in a film of water to
reach and then fertilize the female gamete (egg). In other words, the sperm has to swim through the outside
environment to reach the egg. In other types of land plants, the
male gamete is not flagellated (it cannot swim), and it is
delivered directly to the female gamete location in a form of
internal fertilization.

Heterospory. Mosses and most ferns produce spores of the

same size that are indistinguishable; therefore, they are
generally referred to as homosporous. Seed plants are
heterosporous because they produce two types of spores:
megaspores and microspores. Megaspores are large and
develop into female gametophytes. Microspores are small and
develop into male gametophytes. We will cover the details and
implications of heterospory in the lab on seed plants.

Asexual reproduction in plants

Asexual reproduction plays a very
important role in plants. Asexual
reproduction produces new individuals
without fertilization (i.e., no fusion of
gametes). The offspring produced by
asexual reproduction are genetically
identical to the parent plant. For
example, in some plants, a new
individual can grow from underground
buds produced from horizontal stems, or
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
from roots some distance from the parent plant. Other plants have evolved specialized structures for asexual
reproduction called asexual propagules. Propagules are capable of dispersing away from the parent plant
and forming a new individual.

Terrestrialization
Overview
Terrestrialization is the process whereby plants adapted to life on dry land. The world we know today would not exist if
green plants had not made the transition from living in water to living on land. Plant terrestrialization was a crucial step
in generating biodiversity. The first organisms to live and reproduce on land were probably plants (along with mutualistic
fungi), which enabled other life forms to survive on land by providing shelter, oxygen, and food. The early land plants
increased the amount of oxygen in the atmosphere and helped break down rock into soil.

Terrestrialization: adaptation to life on land & in air

The process of terrestrialization involved adaptation to life on land and in air as opposed to water. The plants that were
able to survive and reproduce on land must have also been able to resist or tolerate desiccation (drying out) in dry air.
The sequence of events leading to the invasion of land by plants likely started when characteristics favorable for survival
in freshwater habitats evolved in algae adapted to marine (salt-water) habitats. The process of terrestrialization
continued as some freshwater green algae adapted to life on land. The freshwater algae likely inhabited the shallow
water around the edges of lakes or ponds where they would have experienced occasional desiccation when the water
level dropped. Traits that allowed these freshwater algae to survive desiccation were likely beneficial to their
descendants as they adapted to life on dry land. Charophytes exhibit some of these desiccation-resistance
characteristics. For example, the zygote is retained within a protective oogonium and is coated with a durable covering
that prevents it from desiccating.

Figure 25. Hypothesized transition from an aquatic freshwater ancestor to land plants (bryophytes, vascular and seed plants).

Land plants or embryophytes

The taxonomic name for land plants, embryophytes, highlights their capacity for nurturing the young sporophyte (the
embryo) within the tissues of the parent gametophyte.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
The first plants to be completely terrestrial were probably physiologically similar to modern bryophytes (mosses,
liverworts, and hornworts). Bryophytes (which are non-vascular plants) are a possibly paraphyletic group containing all
land plants that are not the vascular plants. (We still don't know how mosses, liverworts, hornworts, and vascular plants
are all related and which group(s) is/are sister to the vascular plants!) Vascular plants possess a vascular system made up
of tissues that allow water and the products of photosynthesis (i.e., carbon-based food) to flow through the body of the
plant. These tissues give the plants rigidity and strength, allowing them to resist gravity (in this way, the vascular tissues
are analogous to the vertebrate bony skeleton). Without the evolution of a vascular system, land plants would be
restricted to moist habitats (and there never would have been tall trees). Another important step in the adaptation of
plants to land was the evolution of seeds in the group of vascular plants known as seed plants (gymnosperms and
angiosperms). This was especially important in allowing plants to reproduce in very dry ecosystems (this will be covered
in detail in the next lab).

We will now focus on some of the key characteristics that make plant life on land possible.

Important adaptations in land plants

● Archegonia and antheridia. Archegonia and antheridia are structures known collectively as gametangia. They
are the structures in which gametes are produced in the land plants. Both of these types of gametangia surround
and protect the developing gamete(s) inside. Archegonia are structures in which eggs are produced. Antheridia
are structures that produce and release sperm. Fertilization occurs when a sperm reaches and fuses with the egg
in an archegonium, producing a single-celled zygote. The zygote undergoes cell division and develops into a
multicellular embryo and eventually into an adult sporophyte. While it is developing, the embryo is nourished
and protected from desiccation in the archegonium. Although superficially similar to the oogonia and antheridia
you learned about in Chara, the archegonia and antheridia of land plants are different tissues.

Figure 26. Transverse section of an

archegonium (left) and an antheridium
(right) on the gametophytes of a moss. Look
for the egg in the archegonium. The
antheridium consists of a sterile jacket of
cells surrounding a mass of cells that
produce the sperm.

● Dependent and protected

multicellular embryos. As
described above, the embryos of land plants are retained within the tissues of the gametophyte. By retaining the
embryo, the parent provides shelter from the environment (especially protection from desiccation) and provides
nutrients through
specialized transfer cells
surrounding the
archegonia. This is
analogous to the transfer
of nutrition to the embryo
through the placenta in
most mammals.
Figure 27. Moss archegonium
(left). Cross-sections of
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
archegonia with and without sperm in Physcomitrium (right). (Koshimizu, S. et al.)

● Production of desiccation-resistant spores in sporangia. Sporophytes have specialized structures called

sporangia that contain sporocytes, the diploid cells which eventually undergo meiosis to generate haploid
spores. Land plant sporangia provide special protection to sporocytes and to the developing spores, much as
gametangia provide protection to developing gametes. The spores are covered by sporopollenin, a substance
that makes spores tough and resistant to desiccation.

● Cuticle. The cuticle is a hydrophobic (water-repellant) layer that covers the body of the sporophyte in the
majority of land plants. The cuticle limits water loss from the plant, provides defense against pathogens, and
provides protection from harmful ultraviolet light from the sun. The cuticle creates some challenges for the plant
as well, because it limits the diffusion of carbon dioxide into the plant. Carbon dioxide is necessary for
photosynthesis. Diffusion of carbon dioxide into land plants is enabled by the presence of stomata, tiny openings
in the cuticle that provide microscopic avenues for gas diffusion.
Figure 28. left: An image of a
rosemary leaf cross-section
taken with a scanning electron
microscope. Note the thin waxy
cuticle in yellow on the upper
surface of the leaf. center: Rain
droplets on the surface of a leaf.
Figure 29 (right). Micrograph of stomata, the openings on leaf surfaces that allow gas exchange.

Mosses
Mosses as representation of bryophytes
Bryophytes, with more than 20,000 species worldwide, represent the most ancient lineages of land plants. Today, they
are represented by three lineages: liverworts, hornworts, and mosses. Bryophytes are particularly common and diverse
in temperate and tropical forests where water is abundant, but also exist in less favorable environments such as deserts.

Bryophytes are the surviving lineages from the extraordinary radiation of land plants that occurred in the Devonian
period (~400 million years ago), and exhibit many ancestral characteristics of land plants. As a result, bryophytes are key
to understanding how the first land plants looked, how plants adapted to the hostile land environment from a freshwater
habitat, and how the different groups of land plants are related to each other.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley

Figure 30. Phylogeny showing one configuration of the hypothesized relationships among major groups of land plants. Frangedakis et
al. 2020. The hornworts: morphology, evolution, and development. New Phytologist.

Mosses are the most familiar and diverse bryophytes. However, there are some organisms commonly referred to as
“mosses” that are not actually mosses, such as club mosses (which are more closely related to ferns) or Spanish moss (a
flowering plant). We will focus on mosses as representatives of bryophytes.

How do mosses differ from other land plants?

Mosses differ from other land plants in several aspects of their physiology, ecology, and evolution.

● Dominant gametophyte. The main green part of a moss that is easily visible to the naked eye is the haploid
gametophyte. The gametophyte body is only a few cells thick. The leaves generally possess only one layer of cells
and often lack a cuticle (and therefore dry out quickly). Because the moss gametophyte lives longer and is larger
than the sporophyte, we say that the gametophyte is dominant. Although some mosses produce both female
and male gametophytes, mosses are not heterosporous (i.e., they do not produce two distinct types of spore).

● Dependent sporophyte. Moss sporophytes are dependent on the gametophytes. The sporophytes remain
attached to the parental gametophytes for the entirety of their entire short lives. The gametophyte provides
carbon-based food (photosynthate), amino acids, and water to the sporophyte. Moss sporophytes are among the
simplest and smallest sporophytes in land plants. The sporophyte consists of a stalk bearing a terminal
sporangium that encloses (and eventually releases) the spores.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley

Figure 31 (left). This figure shows the dependent sporophyte attached to the gametophyte in moss, Buxbaumia viridis.

Figure 32 (center). Single sporangium (capsule) showing a specialized peristome that realizes spores according to changes in the
environment in moss Oxyrrhynchium speciosum.

Figure 33 (right). Morphological characteristics of the moss Syntrichia. A: Cross section of a moss leaf showing the midrib in the
center and the one cell layer of thickness of the leaf. B: Leaf top view. C: Hydrated clump of a moss on rock substrate. D. Completely
dry moss stem (gametophyte) that can resurrect after applying a few drops of water.

● Small size and utilization of microhabitats. Mosses have an intimate relationship with their surroundings. They
lack the vascular tissue that moves water and photosynthate (the product of photosynthesis) through the bodies
of vascular plants. With their small size and lack of roots, mosses can absorb water directly from the
environment through their entire body. Mosses require specific microhabitats and substrates. For example, they
can live in small cracks in rocks, or on the bark of trees, as long as they can absorb adequate water and inorganic
nutrients and receive sunlight.
● Desiccation tolerance. Mosses can easily recover after becoming completely desiccated for a long period of
time. They can dehydrate (without dying) when water is scarce, then rehydrate and reactivate their cells when
moisture is available again.
● Dependence on water for sexual reproduction. In mosses, sperm are produced in antheridia and must swim
through the outside environment to reach an egg in an archegonium. This is only possible if enough moisture is
present through which the sperm can swim. In this way, mosses are similar to their algae relatives.

Figure 34. Sexual reproduction requires that sperm (produced in antheridia) swim to the eggs located in the archegonia.

Figure 35 (right). Haplodiplontic life cycle of moss. Note the parts of the
life cycle where water is required and where changes in ploidy take
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
place. Note also that although some mosses produce both female and male gametophytes, mosses are not truly heterosporous–they
do not produce two distinct types of spore.

Ecological benefits: biocrusts and peatlands

In addition to having unique adaptations, mosses also provide major ecological and economic benefits. In many
ecosystems where the climate is too dry or too cold for dense vegetation to grow, mosses are members of biological
crusts, or biocrusts. Biocrusts are vital but fragile systems that cover the surface of the soil. This living “skin” of the earth
is an interconnected system of cyanobacteria, algae, lichens, and mosses. Biocrusts have many important functions in
these ecosystems: they protect soil from erosion, contribute to and recycle nutrients in the soil, and make it easier for
soil to absorb and retain water. Biocrusts are easily damaged by the activity of humans and livestock, and once the crusts
are damaged, it may take centuries for the soil to recover.

Figure 36 (left). Biocrust growing in Canyonlands National Park on the Colorado Plateau. Source: Bill Bowman
Figure 37 (right). The living skin of the Earth-mosses, lichens and cyanobacteria form a complex layer on the soil surface.

Another example of the importance of mosses comes from a single

genus of moss, Sphagnum. Sphagnum is often a major component of
deposits of partially decayed organic material known as peat. Peat is
commonly found in high latitude boreal regions. The low
temperature, pH, and oxygen level in peatlands slow the decay of the
moss. For that reason, their remains can be preserved for thousands
of years. Currently, peatlands cover almost 3% of the Earth’s land
surface and contain 30% of global soil carbon (400 billion tons of
organic carbon). Peatlands play an important role in stabilizing
atmospheric carbon dioxide concentrations. Today, peat is still
harvested for fuel in a few countries such as Ireland and Canada. In
the past, the moss Sphagnum was used to make diapers and naturally antiseptic packing material for wounds.
Figure 38 (above). Someone cutting peat from a deposit.
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
Terrestrialization Lab - Key Terms

The following list of terms will help you identify the main concepts that you need to understand after this lab.
Knowing the definition of each term is a good start, but the challenge is to be able to make connections among
the terms through the life cycles of the different organisms we are studying.

Autotroph vs. heterotroph

Photoautotroph
Cellular organization: Unicellular, multicellular, colonial
Prokaryotes vs. eukaryotes
Organelle
Endosymbiosis
Mitosis
Meiosis
Haploid vs. Diploid
Gametes: egg and sperm
Fertilization
Zygote
Embryo
Spore
Sporangium (pl. sporangia)
Sporophyte
Gametophyte
Isomorphic
Dominant generation
Independent / dependent generation
Gametangia: Antheridium (pl. antheridia) and Archegonium (pl. archegonia)
Oogonium
Motile gametes
Haplodiplontic life cycle (also referred to as alternation of generations)
Haplontic life cycle
Embryophyte (land plant)
 Copyright © Fall 2024, Biology 1B, University of California Berkeley
Terrestrialization
Vascular tissue
Desiccation tolerance
Cuticle
Sorus (pl. sori)
Following this lab you should be familiar with these life cycles:
● The general haplodiplontic life cycle
● The life of Ulva sp.
● The life cycle of Chara sp.
● The moss life cycle