EMBRYOPHYTES

OVERVIEW

- EMBRYOPHYTES: EVOLUTION, HABITAT, CHARACTERISTICS, LIFE CYCLE,

CLASSIFICATION, AND PHYLOGENY.
 Turichire & Bowler (2011)

Eukaryote phylogenetic tree derived from different molecular phylogenetic and ultrastructural studies (adapted from
Baldauf, 2008). Stars indicate algal species for which genome sequences or expressed sequence tags (ESTs) are available,
and images from the representative species, Cyanophora paradoxa for glaucophytes, Micromonas for prasinophytes,
Chroomonas for cryptophytes, Emiliania huxleyi for haptophytes, Ectocarpus siliculosus for brown algae and
Phaeodactylum tricornutum for diatoms. Numbers 1–4 represent the different super groups of eukaryotes. Red arrows
point to the plant, animal and fungal kingdoms.
 Reconstruction of an ancient aquatic–
terrestrial landscape, with the earliest
multicellular land plants, adapted from a
drawing of Z. Burian
(c. 1945) (a). Comparison between the
diplobiontic and haplobiontic life cycles with
some representative botanical examples
(embryophytes
and two aquatic charophytes, Coleochaete
and Chara, respectively) (b). In the
diplobiontic life cycle, the haploid and diploid
phases
(gametophyte, G, and sporophyte, S,
respectively) are multicellular. In the
haplobiontic-haploid life cycle, only the
haploid phase is multicellular.
The haplobiontic-diploid life cycle (in which
the diploid phase exclusively expresses
multicellularity, such as in our species and
other animals)
is not shown. n = number of chromosomes
per haploid cell.
Niklas & Kutschera (2010)
 Niklas & Kutschera (2010)

Concept map with three major themes (axes) underlying a sequential exposition (steps 1–6)
of the evolution of the land plant life cycle. H-d, haplobiontic-diploid; H-h, haplobiontic-
haploid. The aquatic–terrestrial axis is illustrated in Fig. 1a. See text for details.
 Niklas & Kutschera (2010)

Phylogenetic relationships among the major green plant lineages based on an

analysis of DNA sequence data. Dichotomous branches shown as broken lines
indicate weakly supported portions of the tree. The phyletic position of the
Mesostigmatophyceae is particularly problematic (denoted by ‘?’). The distributions
of different habitat preferences, body plans and life cycles are indicated by symbols
(see lower box). Adapted from Lewis & McCourt (2004).
 Phylogenetic relationships among the main
lineages of green plants. The tree topology is a
composite of accepted relationships based on
molecular phylogenetic evidence [1, 9–11, 22, 23,
35, 36, 56, 71, 78]. Uncertain phylogenetic
relationships are indicated by polytomies. The
divergence times are rough approximations based
on the fossil record and molecular clock estimates
[2–4, 13, 14]. These age estimates should be
interpreted with care as different molecular clock
studies have shown variation in divergence times
between major green plant lineages. Drawings
illustrate representatives of each lineage: (1)
Acetabularia, (2) Pediastrum, (3) Chlorella, (4)
Tetraselmis, (5) Picocystis, (6) Ostreococcus, (7)
Micromonas, (8) Crustomastix, (9) Monomastix,
(10) Pyramimonas, (11) Pycnococcus, (12)
Pseudoscourfieldia, (13) Nephroselmis, (14)
Prasinococcus, (15) Verdigellas (a: general habit,
b: individual cells in a gelatinous matrix), (16)
Mesostigma, (17) Chlorokybus, (18)
Klebsormidium, (19) Chara, (20) Xanthidium, (21)
Chaetosphaeridium, (22) Coleochaete, (23)
Leliaert et al. (2011) Ranunculus.
Schematic representation of the
evolutionary relationships and
divergence times for the red, green,
glaucophyte, and chromist algae.
These photosynthetic groups are
outgroup-rooted with the
Opisthokonta which putatively
ancestrally lacked a plastid. The
branches on which the
cyanobacterial (CB) primary and red
algal chromist secondary
endosymbioses occurred are
shown.
Leliaert et al. (2012)
 ADAPTACIONES A LA VIDA TERRESTRE

♦ Cambio en el ciclo de vida.

♦ Aparece la cutícula protegiendo al esporofito.

♦ Aparece la esporopolenina formando una gruesa pared en la espora.

♦ El intercambio de gases con el ambiente.

♦ Aparecen los flavonoides.

♦ Aparece el sistema de oxidación del glicolato.

♦ Se produce la interacción entre las plantas terrestres y los hongos.

 ¿Qué ventaja adaptativa encontraron los clados más modernos de embriofitas,
que los volvieron dominantes de la superficie terrestre?

I. El esporofito, al tener dos juegos de cromosomas en lugar de uno, es menos

vulnerable ante las mutaciones genéticas, lo cual como consecuencia aumenta la
diversidad de genes y por lo tanto las probabilidades de estar adaptado cuando
cambia el ambiente.

II. El esporofito no tiene limitantes ecológicos para crecer en altura, ya que no necesita
ningún tipo de interacción para producir las esporas y liberarlas. En cambio el
gametofito, productor de gametos masculinas y femeninas, debe tener una
constitución que le asegure que las gametos masculinas puedan nadar hasta las
femeninas, para que se produzca la fecundación y se reinicie el ciclo. Esto limitaría
en gran medida la exploración de diferentes morfologías en el gametofito, que debe
estar ligado al suelo para asegurar la fecundación y por lo tanto la supervivencia de
la especie (Judd et al., 2002).
Evolution of apical growth. The common
ancestor of embryophytes and their
closest relatives display apical growth in
the gametophyte generation (Charales
lack an alternation of generations; the
only diploid phase is the zygote). Apical
growth in the sporophytes of mosses
and polysporangiophytes may or may
not be homologous (in terms of a
developmental process expressed in the
sporophyte). Loss of apical growth in the
gametophyte generation is associated
with transitions to heterospory.

Friedman et al. (2004)

 Friedman et al. (2004)

(A) In the absence of

integration of fossil taxa into a
phylogeny, it might be
concluded that the roots of all
extant plants are homologous,
but this is an artifact because
key fossil taxa appear to lack
roots.
(B) When key fossil taxa are
integrated into the phylogeny for
analysis of root evolution, it is
apparent that there have been two
evolutionary origins of roots. One
origin is in the Lycophyta, perhaps
from telomic systems that grow into
substrate, as found in
Zosterophyllum. Another
independent origin is observed in
the common ancestor of extant
euphyllophytes.

Friedman et al. (2004)

 Evolution of leaves. A minimum of
three evolutionary origins of leaves
are inferred in the gametophyte
generation. One origin is among
complex thalloid liverworts,
another in simple thalloid
liverworts, and a third in the
common ancestor of extant
mosses. Three evolutionary origins
are also inferred in the sporophyte
generation. One origin is in
Lycopsida, another in the
moniliformopses, and a third in the
common ancestor of seed plants
and Archaeopteridales.
Friedman et al. (2004)
BRYOPHYTES
HORNWORTS
MOSSES
LIVERWORTS
CLASSIFICATION
LYCOPHYTA
1. It is the oldest extant (living) vascular plant division
at around 410 million years old.

2. These species reproduce by shedding spores and

have macroscopic alternation of generations, although
some are homosporous while others
are heterosporous.

3. Members of Lycopodiophyta bear a protostele, and

the sporophyte generation is dominant. They have
microphylls, leaves that have only a single vascular
trace (vein) rather than the much more
complex megaphylls found in ferns and seed plants.
CLASSIFICATION

•Class Lycopodiopsida – clubmosses and firmosses

•Class Isoetopsida – quillworts, scale trees, and spikemosses
•Class † Zosterophyllopsida – extinct zosterophylls.

LYCOPODIUM ISOETES SELAGINELLA

MONILOPHYTA
Equisetaceae
GIMNOSPERM
Ginkgo biloba
CYCADACEAE
PINACEAE
ANGIOSPERM
MONOCOTS DICOTS
MONOCOTS

DICOTS
DICOTS
MONOCOTS DICOTS
MONOCOTS
DICOTS
 PHYLOGENY

Cole & Hilger (2013)