BIOLOGY DEPARTMENT, DE LA SALLE UNIVERSITY

SYSTEMATICS LABORATORY (LBYBI12) TERM 3 AY 2019-2020

Exercise 7

BRYOPHYTES
(Mosses, Liverworts and Hornworts)

Introduction

Bryophytes represent the most primitive extant members of the Kingdom Plantae. They lack a specialized vascular or conducting
system to transport water, dissolved nutrients and essential inorganic substances throughout the plant body. In the absence of such
vascular system, they rely on diffusion and osmosis to obtain their needed nourishments. This restricts their size and makes them
dependent on moist environment for active growth and development.

Among the bryophytes’ major contrast from their green algal ancestors is the development of a jacket of sterile cells that surround
the archegonia and antheridia, the structures that produce the egg and sperm, respectively. These gametes are formed in the large
photosynthetic gametophytes (n) that are attached to the substrate by root-like structures called rhizoids. Such gametophytes are
dorsoventrally flattened and bilaterally symmetrical in the liverworts and hornworts but are erect and radially symmetrical in mosses.
The gametophytes support the sporophytes (2n) that produce similar type spores, a condition called homospory.

All bryophytes are classified under three distinct divisions: Division Bryophyta (trad. Musci; mosses), Division Hepatophyta (trad.
Hepaticae; liverworts) and Division Anthocerophyta (trad. Anthocerotae; hornworts). To date, about 16,000 species of mosses,
liverworts and hornworts have been described in science. They are all non-vascular and share similar life cycles but also differ in many
ways. In this exercise, representatives of these plant divisions will be examined.

Objectives
1. To distinguish the bryophytes based on their morphological and reproductive features.
2. To identify the relationships of the members of the bryophytes within its members and to other land plants
3. To explore the diversity, economic and ecological importance of bryophytes.

Materials
Images of live specimens
Photomicrographs of whole mounts and specimen sections

Procedure
A. Bryophyte Phylogenetics

Figure 1 shows phylogenetic relationships of the

classical Bryophytes within its members and the higher
forms of plants. The bars on the lineages indicate the
synapomorphies (shared-derived features) of all land
plants (including the Bryophytes), and the autapomorphic
features (derived or unique features; diagnostic
characters) per group.

Based on the diagram, answer the following questions:

1. Name the synapomorphies the made the Bryophytes Figure 1. Most recent phylogenetic relationships within the Embryophyta
inclusive of the Land Plants. based both morphological and molecular data sensu Puttick et al. 2018.
Illustration taken from Simpson 2019.
2. Identify the morphological feature that separated the
Bryophytes from the rest of the members or the Land plants
3. Identify the phylogenetic relationship of the Bryophytes. Is it resolved or not? Support your answer.
4. On the phylogenetic positions depicted in the given tree, would you consider liverworts more primitive than the mosses? Why or
why not?
 BIOLOGY DEPARTMENT, DE LA SALLE UNIVERSITY
SYSTEMATICS LABORATORY (LBYBI12) TERM 3 AY 2019-2020

B. Division Bryophyta (Mosses)

Mosses consist of close to 10,000 species that grow in colonies on rocks, soil or tree trunks. The individual moss plant
has a leafy axis consisting of a caulidium ("stem") and radially arranged phyllidia ("leaves"). The phyllidia have no mesophyll tissue,
stomata or veins. They are generally sessile and are merely one cell thick except at the costa, the structure that runs through the
middle of the phyllidium. The cells in these phyllidia usually contain numerous lens-shaped chloroplasts except at the costa. The
phyllidia of Sphagnum (peatmoss) however, have huge, transparent water-storage hyaline cells that adapt them to water
absorption and storage. Small green photosynhetic cells are sandwiched between these large hyaline cells. Few mosses can grow
to over a foot long. They develop specialized water-conducting cells or hydroids and food -conducting cells or leptoids.

1. Photomicrographs of sample mosses will be provided as reference. Draw and label the parts: gametophyte with caulidia
and phyllidia and the sporophyte (foot, seta and capsule).
2. Examine an image of the tip of a leafy plant. If the tip has a flat "flowerlike" cluster of leaves, the plant is a male
gametophyte and the male reproductive organs are located in this flat circlet of leaves. If the leaf arrangement at the tip
is no different from the rest of the stem, the plant is a female gametophyte and the female reproductive organs occur
hidden among the leaves at the tip. These leafy gametophytes develop at first from the filamentous gametophyte called
protonema.
a. Based on this information, acquire or draw the images of the following stages of development (sporophyte and
gametophyte) and then arrange them in a complete “image” life cycle diagram. Place the labels under each image,
indicating their ploidy stages (haploid or diploid). On the arrows, indicate the events taking place (mitosis, meiosis)
3. Examine the provided images of Mnium and Sphagnum. Draw and label the following parts: protonema (with buds),
antheridiophore (with antheridia, sterile jacket, sperm cells, paraphyses – sterile filaments), archegoniophore (with
archegonia, egg, venter, neck, paraphyses), capsule (calyptra, operculum, peristome, columella, spores, annulus).
C. Division Hepatophyta (Liverworts)
Liverworts comprise of about 6,000 species that often form flattened, lobed structures called thalli (sing. thallus), i.e., a
plant body not differentiated into stems, leaves, and roots. They are so named “liverworts” because the lobes of their thallus
resemble the lobes of an animal liver. Like mosses, liverworts have dominant photosynthetic gametophytes. Based on the
appearance of these gametophytes, liverworts are classified as thallose liverwort (Oder Marchantiales) and leafy liverwort (Order
Jungermanniales).

1. Examine a photomicrograph or an actual image of Marchantia. Note the characteristic dichotomous (Y-shape) branching
of the gametophyte thallus. Examine further a magnified image of the thallus. At the edges, locate the slender, hair-like
structures known as the rhizoids. Associated with the rhizoids are long, flattened structures, the scales. Examine the
dorsal surface and observe the minute “pores”. In live and/or actual specimens, these pores are visible with a hand lens.
2. Asexual reproduction
Older gametophytes of Marchantia form asexual reproductive structures known as the gemmae (sing. gemma) cups.
These small cups contain a number of multicellular structures called gemmae that are attached by a short stalk to the
base of the cup. When mature, the gemmae becomes detached and are easily dispersed, germinating to form a new
thallus in suitable conditions. Draw a botanical sketch of a thallus with gemmae cups (with gemmae).
3. Sexual reproductions
The sexual reproductive organs of the Marchantia are borne on special upright branches. Each branch has a stalk and a
terminal disc. The male and female gametophytes are distinguished by the shape of their terminal discs. Many species
of Marchantia are dioecious; that is, there are separate and therefore two kinds of reproductive branches
a. Examine the provided image of the gametophyte specimens of Marchantia. The male gametophyte bears an
antheridial branch called antheridiophore, whereas the female gametophyte bears an archgonial branch called
archegoniophore. Note the difference in the shape of the terminal disc. Archegoniophores have deeper lobes
compared with the umbrella-like form of the antheridiophore.
b. Examine the microscopic images of both the antheridiophore and archegoniophore. In the antheridiophore,
look for the flask-shaped antheridia on the upper surface of the disk. The antheridium is made up of sterile
jacket of cells, spermatogenous cells, and stalk cells. In mature ones, sperms are extruded in a mucilaginous
mass. In the archegoniophore, look for the flask-shaped archegonia which are situated in rows on the under
surface of the disk. Each row is separated from its neighbour by an outgrowth of tissue called perichaetium.
Study in detail the different parts of the archegonium.
i. Neck – an elongated portion which leads to an opening.
 BIOLOGY DEPARTMENT, DE LA SALLE UNIVERSITY
SYSTEMATICS LABORATORY (LBYBI12) TERM 3 AY 2019-2020

ii. Venter – an enlarged basal portion of the archegonium consisting of ventral canal cells, a haploid egg,
and the venter wall cells. Draw and label the parts of the archegoniophore and the antheridiophore.
4. The Sporophyte
The embryo of the young sporophyte is formed as a result of the fusion of sperm and egg nuclei. It develops within the
venter of the archegonium. The mature sporophyte consists of the following parts: foot, seta (pl. setae), capsule
(=sporangium), spores, and elaters.

D. Division Anthocerophyta (Hornworts)

Hornworts (ca. 100 species) are a group of small, inconspicuous thalloid plants that grow on moist soil, hidden under
grasses and other herbs. Hornworts superficially resemble thalloid liverworts, the gametophytes of many species being ribbon-
shaped and thin, without a distinct stem of leaves. However, hornworts lack oil bodies contrary to the liverworts. As in all other
embryophytes, an alternation of heteromorphic generations occurs, with the sporophyte, depending on the larger
photosynthetically active gametophyte.
1. The Gametophyte
The hornwort gametophyte is the simplest of the bryophytes. They are small, green discoid thallus with little internal
differentiation of vegetative tissues. They are slightly lobed, with numerous rhizoids growing from the lower surface. The
antheridia are similar in structure to that of the hepatics. They are located in roofed chambers in the upper portion of
the thallus. The archegonia are embedded within the thallus just like the antheridia. As in the case of mosses and
liverworts, some species of hornworts have unisexual plants whereas other species are bisexual.
2. The Sporophyte
The sporophyte is in striking contrast to those of the liverworts. The sub-epidermal cells contain chloroplasts while the
epidermis has stomata. A foot embedded in the thallus serves as an absorbing organ. The sporangium is an upright,
elongated structure that grows like a horn from the gametophyte and contains spores that mature in progression from
top down.
3. Examine the thallus of Anthoceros with sporophyte under the dissecting microscope. Draw and label the parts.

References:
Balbach MK, LC Bliss. 1991. A Laboratory Manual for Botany (7th ed.) Brooks/Cole, Thomson Learning pp. 277-292.
Berg L. 2012. Introduction to Botany. Cengage Learning Asia Pte Ltd. Pp. 470-480.
Puttick et al. 2018. In Simpson MG. 2019. Plant Systematics (3rd Ed.). Elsevier (Academic Press). Unit II, Chapter 3. (Digital Copy)
Reece JB, LA Urry, ML Cain, SA Wasserman, PV Minorsky, RB Jackson. 2011 Campbell Biology, Global Edition (9th). Pearson Education, Inc.
Simpson MG. 2019. Plant Systematics (3rd Ed.). Elsevier (Academic Press). Unit II, Chapter 3. (Digital Copy)

Revised by Dr. Emelina Mandia (28.May.2019)

Modified by Bismark Oliver Lemana (24.June.2020)