ICBI 231 Plant Biology

The origin of algae

and evolution of
embryophytes

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Outline:

▪ Origin of eukaryotic cells

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▪ Endosymbiosis theory
▪ Mitochondria
▪ Plastids

▪ Types of cytokinesis in algae and plants

▪ Origin of embryophytes (terrestrial plants)
▪ Green algae

Figure 20-2 (A) Euglenoid algae. (B) A dinoflagellate. (C) Centric diatoms. (D) Golden-
brown algae. (E) Yellow-green algae. (F) Green algae. (G) Brown algae. (H) Red algae.
Fun facts about Algae

▪ Many algae in lakes and streams are mistakenly called moss

▪ Algae live in soil, more live in freshwater, and many inhibit oceans, but none lives at

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water depths where sunlight does not penetrate.
▪ A photosynthetic cyanobacterium evolved into a chloroplast after entering eukaryotic
cells .
▪ This line gave rise to algae, including green algae, which in turn produced true plants,
embryophytes.
Origin of Eukaryotic Cells

▪ The differences between prokaryotes and eukaryotes are metabolic processes and
cellular organization.

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▪ DNA structure: prokaryotic DNA is circular, Eukaryote DNA is linear and bound to
protein
▪ Nuclear structure and division: prokaryotes lack a nucleus, DNA lies in the cytoplasm.
▪ Organelles: Prokaryotes lack membrane-bound organelles.
Origin of Mitochondria and Plastids

▪ Until the early 1970s, the autogenous theory

was accepted, in which organisms gradually

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became more complex.
▪ In 1905, K. C. Mereschkowsky speculated that
plastids and mitochondria might be prokaryotes
living inside eukaryotic cells (endosymbiont
theory).
▪ By the late 1960s, plastids and mitochondria were
discovered to have their own DNA and
ribosomes, both having prokaryotic features.
▪ DNA sequence data indicate mitochondria are
most closely related to proteobacteria. FIGURE 20-5 The endosymbiont theory
postulates that a prokaryote.
▪ Endosymbiosis occurred ~ 2 million years ago.
Origin of Mitochondria and Plastids

▪ Chloroplasts likely evolved when a eukaryotic

cell containing mitochondria engulfed a

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photosynthetic cyanobacteria cell (SF Fig. 2.4
Part 5). This is also called primary
endosymbiosis.
▪ have two sets of cell membranes surrounding
them: one from the host cell and one from the
endosymbiont.

▪ Secondary endosymbiosis occurs when a

eukaryotic cell engulfs a cell that has already
undergone primary endosymbiosis.
▪ have more than two sets of membranes
surrounding the chloroplasts.
Types of Cytokinesis

▪ Several types of cytokinesis occur in algae.

▪ In almost all algae with walls, cell division is similar to

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that of plants by use of a phragmoplast.
▪ In some green algae, cytokinesis occurs by a
phycoplast, with microtubules oriented parallel to the
plane where the new wall will form, which is
perpendicular to the orientation of the spindle.
Origin of embryophytes

▪ Early green algae organized complex

differentiated multicellular bodies.

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▪ Some moved onto land—the ancestors of
embryophytes, true plants.
▪ specifically charophytes, during the
Ordovician period (~500 million years ago).

▪ The transition to land was driven by

selective pressures such as increased
sunlight and carbon dioxide availability.

FIGURE 20-2 (F) Green algae.

Origin of Embryophytes

▪ Alternation of generation: a life

cycle pattern alternating between a

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sexual phase and an asexual
phase:
▪ Found in both algae and plants

▪ Monobiontic life cycle – a single

independent generation. i.e. there
is no alternation of generations.
▪ Simplest life cycle

FIGURE 20-14 In monobiontic life cycles, only one generation—one phase—is capable
of undergoing mitosis.
Green Algae: Life Cycles

▪ In a dibiontic species in which both stages are multicellular, the gametophyte

(haploid phase) and sporophyte (diploid phase) may resemble each other strongly;

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this is alternation of isomorphic generations.
▪ If the two phases are very different in appearance and construction, the life cycle is
an alternation of heteromorphic generations.

FIGURE 20-15 Dibiontic life cycles involve an alternation of either isomorphic (A) or heteromorphic (B) generations.
Green Algae: Life Cycles

▪ During the earliest stages of the evolution of sexual

reproduction, gametes were isogamous (identical).

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▪ Anisogamy and oogamy later evolved.

▪ Gametes are produced in gametangia.

(A) Isogametes
▪ Spores are formed in sporangia.

(B) Anisogametes

FIGURE 20-16 (A) Isogametes are identical gametes. Theoretically, they could be either
motile or nonmotile, but in reality, all known isogametes do have flagella. (B) If gametes
have any visible differences but are still similar, they are anisogametes. (C) Oogametes
are bviously different in size, and the megagamete is virtually always nonmotile.
(C) Oogametes
Green Algae: Representative Genera

▪ Filamentous species
▪ Ulothrix species’ life cycle is monobiontic; it involves only

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one free-living multicellular generation, and it is haploid.
▪ Spirogyra is an extremely common fresh-water filamentous
green alga that reproduces through conjugation.

▪ Laminar species
▪ Ulva has a dibiontic life cycle with an alternation of
isomorphic generations.

▪ Coenocytic species © Michael Abbey/Science Source; © Blickwinkel/NaturimBid/Alamy Stock

▪ Derbesia has a dibiontic life cycle with an alternation of Photo

FIGURE 20-19 (A) Individuals of Ulothrix are uniseriate,
heteromorphic generations. unbranched filaments of haploid cells (×80). (B) Cells of
Spirogyra, showing the spiral, band-shaped chloroplasts.

▪ Parenchymatous species
▪ One group, chlorophytes, divides by means of a
phycoplast.
▪ The other group, charophytes, undergo cell division by
means of a phragmoplast, just as plant cells do.
Green algae and embryophytes
▪ An interesting example is Chara, which has a
stem-like body divided into nodes and internodes,

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with whorls of branches arising at internodes.
▪ Reproduction in Chara is significant;
embryophytes have multicellular reproductive
structures with sterile cells.
▪ Charophytes and embryophytes are sister groups
sharing a common ancestor.
▪ As such, they form a monophyletic clade, and
some people have suggested that this clade be
named the streptophytes.
▪ Other suggested names include the
archaeoplastids or primoplantae.
FIGURE 20-22 (A) An individual of Chara, with a
parenchymatous body, microgametangia (spherical),
and megagametangia (oval).
ICBI 231 Plant Biology

Nonvascular Plants:
Mosses, Liverworts,
and Hornworts
Dr. Sunisa meanchaipiboon
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Outline

▪ Characteristics of nonvascular plants

▪ Classification of nonvascular plants

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▪ Division Bryophyta: Mosses
▪ The gametophyte generation
▪ The sporophyte generation
▪ Metabolism and ecology

▪ Division Hepatophyta: Liverworts

▪ The gametophyte generation
▪ The sporophyte generation

▪ Division Anthocerotophyta: Hornwarts

▪ The gametophyte generation
▪ The sporophyte generation
Classification of plants

▪ Plants are traditionally divided into three

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groups.
▪ Neither vascular tissues nor seeds, the
nonvascular plants (often called
“bryophytes”).
▪ Vascular tissue but not seeds, the
vascular cryptogams.
▪ Both vascular tissue and seeds,
spermatophytes.
▪ Beginning at some point earlier than 450
million years ago, certain green algae
known as charophytes began to adapt to FIGURE 21-1 Recently proposed phylogeny of true plants. MYBP = million
years before present
living on land.
The origin of embryophytes

▪ They evolved to survive the occasional drying of their streams, small lakes, and
oceanside mud flats.

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▪ Due to dry land conditions, gamete and spore mother cells needed protection from
dryness.
▪ As a truly terrestrial existence became more successful, the environment became
selective for mutations that produced an upright body that could grow into brighter light.
▪ Production of pollen and seeds eliminates the need for environmental water for
reproduction.
▪ Vascular tissue, especially phloem, also made feasible the evolution of truly heterotrophic
tissues—roots, meristems, and organ primordia.

FIGURE 21-3 Megagametangia of true plants, such as these of a moss.

Characters of Nonvascular Plants

▪ Have multicellular sporangia and gametangia and a

life cycle with an alternation of heteromorphic

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generations.
▪ All mosses and many liverworts have leafy stems that
look remarkably like small versions of flowering
plants.
FIGURE 21-2 In true plants, gametangia and sporangia
▪ Nonvascular plants are almost exclusively terrestrial always have an outermost layer of cells that do not
become gametes or spores.
and have a cuticle over much of their bodies, and
many have stomata.

FIGURE 21-5 The megagametophyte of

Selaginella, which has developed almost
completely within the original wall of the
megaspore.
FIGURE 21-6 The tiny leafy shoots are gametophytes
of a small moss.
Classification of Nonvascular Plants
https://www.brainkart.com/article/Bryophytes_32870/
▪ It is not known how closely related
mosses, liverworts, and hornworts

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are.
▪ They are often treated as three
distinct divisions.
▪ Liverworts, division Hepatophyta
▪ Mosses, division Bryophyta
▪ Hornworts, division
Anthocerotophyta
Division Bryophyta: Gametophyte

▪ Mosses are ubiquitous, occurring in all parts of the world and in

almost every environment. They are perennial and thrive in

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many places within cities.
▪ The gametophyte generation
▪ The leafy stems are technically known as gametophores and
form dense mounds.
▪ All moss stems have leaves, but because they are parts of a FIGURE 21-7 Although mosses occur in hot,
gametophyte, not a sporophyte, they are not homologous with arid areas, they are most abundant in cool,
moist regions.
those of vascular plants.

▪ In some mosses, the innermost cortex is composed of cells

called hydroids that conduct water and dissolved minerals.
▪ Species that have hydroids typically also have leptoids, cells
that resemble sieve cells. © James D. Mauseth

FIGURE 21-4 (A) This rock is covered with a moss

(Grimmia laevigata, the gray-green material) and
lichens (the blue-gray material). (B) Is somebody
mixed up? Mosses, liverworts, and cacti side by
side.
Division Bryophyta: Gametophyte

▪ The majority of mosses lack hydroids and leptoids;

water is conducted along the exterior of their stems

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by capillary action.
▪ At the base of the stem are rhizoids, which anchor
the stem but do not absorb water or minerals.
▪ Gametophyte generation forms when a spore
germinates and produces a protonema filament that
gives rise to multiple gametophores.

FIGURE 21-15 A moss spore has germinated and grown

into these filaments of cells, which make up a protonema;
later some cells will divide into apical meristems that will
grow into gametophores.
Division Bryophyta: Gametophyte

▪ The gametophore at some point produces gametangia; antheridia produce

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sperm and archegonia produce eggs.
▪ Antheridia and archegonia occur on the same gametophore in bisexual species
(Funaria, Pottia), whereas other species have both male and female gametophores
(Barbula, Polytrichum, Rhacomitrium).
▪ Secretion of sucrose from the archegonia guides sperm as they swim toward the
archegonia and then to the egg, where one sperm cell effects fertilization.
Division Bryophyta: Sporophyte

▪ Moss gametophytes are both large and

photosynthetic, and they support the

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sporophyte throughout its entire life.
▪ The moss sporophyte is never an
independent, free-living plant.
▪ All moss sporophytes form from the zygote
and have three basic components.
▪ Foot: the interface with the gametophore.
▪ Capsule: simple sporangium where spores
are produced.
▪ Seta: between the foot and the sporangium.
▪ None is branched or has leaves, bracts, or FIGURE 21-10 The individual shoots of a
moss superficially resemble those of a
FIGURE 21-19 Aspects of a
moss sporophyte.
buds of any kind. flowering plant, having stems, leaves,
nodes and internodes, and even buds.
Division Bryophyta: Sporophyte

▪ Dehiscence of the sporangium is more

complex than opening of the gametangia.

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▪ The apex of the sporangium is a cap-like
lid called the operculum which separates
from the rest of the sporangium.
▪ Peristome teeth respond to humidity,
bending inward and opening the
operculum when the air is dry.
▪ The apex is covered by a calyptera.

© Biophoto Associates/ Science Source

FIGURE 21-19 Aspects of a moss sporophyte.

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FIGURE 21-16 Life cycle of a moss.
Details are given in the text.
Division Bryophyta: Life Cycle
Division Bryophyta: Metabolism and Ecology

▪ The small size and lack of conducting tissues are two critical factors in the
metabolism and ecology of mosses.

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▪ Without vascular tissue, stems and leaves can become desiccated, even while
the rhizoids are in contact with moist soil or tree bark.
▪ Several mechanisms compensate for the inability of mosses to retain water,
such as life in moist habitats.
Division Bryophyta: Mosses

▪ Other mosses are tolerant of desiccation; drying does

not damage them as it does most vascular plants and

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algae.
▪ Desiccated mosses are remarkably resistant to high or low
temperature and to intense ultraviolet (UV).
▪ Many moss species thrive at low temperatures near or
even below 0°C.

▪ Certain mosses can grow on hard, impervious surfaces

because they have no roots that must penetrate the
substrate.

FIGURE 21-21 (D) In dry seasons, plants of

Oxymitra desiccate and fold inward, bringing the
scales over their body, protecting themselves.
They survive months of severe heat and drought.

FIGURE 21-4 (A) This rock is covered with a

moss (Grimmia laevigata, the gray-green
material) and lichens (the blue-gray material).
Division Hepatophyta: Liverworts

▪ Like mosses, liverworts are small plants that

have an alternation of heteromorphic

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generations.
▪ The sporophyte is even less conspicuous than
in mosses and is completely dependent on the
gametophyte.
▪ Hepatic gametophytes are divided into two
basic groups.
▪ Leafy liverworts
▪ Thallose liverworts
Courtesy of Dr. Andrew Spink (www.andrewspink.nl)

▪ The gametophore of leafy liverworts resembles FIGURE 21-21 (A) Leafy liverworts such as this
Lophocolea can be very easily confused with mosses.
that of a moss—thin leaves on a slender stem.
Division Hepatophyta: Gametophyte

▪ Thallose liverworts are not leafy at all but rather flat and
ribbon like or heart shaped and bilaterally symmetrical;

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this shape is called a thallus.
▪ Many cells contain large oil bodies.
▪ Cells are loosely arranged as aerenchyma with large air © Biophoto Associates/Science Source
FIGURE 21-23 Although gametophytes of thallose
chambers that open to the exterior by means of large air liverworts may become very thick (a few
millimeters), they are never solid; rather, they have
pores. numerous chambers, and both carbon dioxide and
water vapor can diffuse through them easily.
▪ Liverwort gametophores may be either bisexual,
producing both antheridia and archegonia, or unisexual,
depending on the species.
▪ Male gametophores produce a stalked, umbrella-shaped
antheridiophore.
▪ Archegoniophores have a set of drooping projections
that droop downward. FIGURE 21-21 (C) Thallose liverworts have rather thick
bodies and are not easily confused with mosses.
 Division Hepatophyta: Liverworts

▪ If sperm cells are carried to the archegoniophore by raindrop splashing, they swim
into the archegonium neck and fertilize the egg.

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▪ The zygote is retained on the gametophore and grows into a small sporophyte.
▪ Most liverwort sporophytes have a foot, seta, and calyptra-covered sporangium.
▪ Within the sporangium, some cells do not undergo meiosis but rather differentiate into
elaters.

FIGURE 21-25 (B) Each archegonium consists of a FIGURE 21-26 Liverwort sporophytes consist of foot,
FIGURE 21-24 (B) Low-magnification view of
long, tubular neck and a slightly swollen base, all only seta, and capsule (sporangium), but no elaborate set of
antheridiophore with antheridia (×8).
one cell thick. teeth as in mosses.
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FIGURE 21-22 Life cycle of
Marchantia. Details are
given in text.
Division Hepatophyta: Life Cycle
 Division Anthocerotophyta: Hornworts

▪ Hornworts are a group of small, inconspicuous thalloid plants that grow on moist
soil, hidden by grasses and other herbs.

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▪ Hornworts superficially resemble thalloid liverworts, but never contain oil bodies.
▪ One of the most striking features is the presence of a single large chloroplast in
each cell as opposed to numerous small plastids present in all other nonalgal plants.

FIGURE 21-9 Hornworts such as

this Phaeoceros are quite rare,
and few people ever see them;
they are easily confused with
liverworts unless the tall “horns”—
sporophytes—are present.
FIGURE 21-29 (A) Hornwort cells are unique among embryophytes: Each has
just one plastid. (B) Cells of privet leaf with many chloroplasts each (360).
Division Anthocerotophyta: Gametophyte
▪ As few as three or four protonema cells are produced before the gametophore phase is
established.

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▪ The gametophyte is parenchymatous, rather succulent but brittle.
▪ Mucilage chambers are invaded by Nostoc cyanobacteria; all hornworts form this symbiosis
and presumably benefit by receiving nitrogen compounds from the Nostoc.
▪ Gametangium development in hornworts is distinctive.
▪ Archegonia are formed from superficial cells, but archegonia do not completely surround the.
▪ After fertilization occurs, the zygote divides transversely.

FIGURE 21-30 (D) The gametangia of hornworts are unique; the antheridia is not surrounded by discrete archegonial cells.
Division Anthocerotophyta: Sporophyte

▪ Similarities between sporophytes of hornworts and those of

mosses or liverworts are not easy to find.

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▪ Hornworts have a foot embedded in gametophore tissue, but
there is no seta or discrete sporangium.
▪ Several attempts have been made to remove the sporophyte
surgically from the gametophyte and grow it in laboratory
conditions, but even though it is chlorophyllous, it dies.

FIGURE 21-31 (A) The base of the sporophyte

resembles a foot embedded in the gametophyte,
and recently transfer cells have been discovered;
therefore, active nutrient transport into the
FIGURE 21-32 A longitudinal sporophyte must be occurring. Just above the foot
section through the sporophyte. is a meristematic region. (B) At a higher level,
equivalent to Figure 21-32C, spores are mature.