Charophytes as the ancestor of land plant

The emergence of land plants or also known as embryophytes approximately

480 million years ago was one of the major events in the history of life on
earth. It led to major changes in the earths environment as for example
reducing the atmospheric CO2 concentration, which consequently lowered
the earths surface temperature, initiated the development of the entire
terrestrial ecosystem and set the stage for evolution of other terrestrial
organisms. There are several groups that are closely linked to the origin and
early evolution of land plants which are charophytes, bryophytes and
pteridophytes.
It is widely accepted that embryophytes evolved from green algae, or more
specifically, from a small but diverse group of green algae known as the
streptophyte algae (charophycean algae). Streptophyte algae and
embryophytes together constitute the division Streptophyta, which likely
split from the Chlorophyta (all other green algae) about 725-1200 MY ago.
Streptophyta and Chlorophyta comprise the Viridiplantae, one of the three
evolutionary lineages derived from the single primary endosymbiosis of a
cyanobacterium and a eukaryotic host cell.
Charophytes are a group of predominantly freshwater green algae from
which land plants are believed to have arisen. They consist of five orders:
Charales, Chlorokybales, Coleochaetales, Klebsormidiales and Zygnematales
(including desmids). The first evidence linking charophytes and land plants
came in the mid-1970s from ultrastructural studies of reproductive cells and
cell division. Since then, numerous lines of evidence have confirmed this
relationship. Phylogenetic analyses of nuclear small subunit (nuSSU) rRNA
sequences have demonstrated that charophytes and land plants form a
monophyletic group, and that this clade, termed Streptophyta, is sister to a
clade composed of all other green algae.
However, there is one central question surrounding the charophytes, that is
which specific lineage is the sister of land plants? Despite many years of
phylogenetic studies, this question has not been resolved and remains
controversial.
To discuss this question, I have a review few journals that propose evidences
either supporting or declining the idea that land plants are closely link with
charophytes.
The first journal that I have study is by

Richard M. Bateman in EARLY EVOLUTION OF LAND PLANTS:

Phylogeny, Physiology, and Ecology of the Primary, Terrestrial
Radiation

Karol K, McCourt R, Cimino M, Delwiche CF (2001) The closest living relatives of land
plants. Science 294: 23512353.
It was nearly a decade ago that Karol et al. concluded after a four-gene, three genome analysis
that, of the charophytes, the Charales constitute the closest living relative to land plants. Another
data analysis supported the same notion and, for a time, this appeared to be a settled matter.
M. Turmel et al. in their structural analyses also revealed that many of the features conserved in
land plant cpDNAs were inherited from their green algal ancestors. The intron content data
predicted that at least 15 of the 21 land plant group II introns were gained early during the
evolution of streptophytes and that a single intron was acquired during the transition from
charophycean green algae to land plants.

The Closest Living Relatives of Land Plants

Kenneth G. Karol, Richard M. McCourt, Matthew T. Cimino,Charles F. Delwiche1
This analysis supports the hypothesis that the land plants are placed phylogenetically within the
Charophyta, identies the Charales (stoneworts) as the closest living relatives of plants, and
shows the Coleochaetales as sister to this Charales/land plant assemblage. The results also
support the unicellular agellate Mesostigma as the earliest branch of the charophyte lineage.
These ndings provide insight into the nature of the ancestor of plants, and have broad
implications for understanding the transition from aquatic green algae to terrestrial plants.
The chloroplast genome sequence of Chara vulgaris sheds new light into the closest green
algal relatives of land plants.
Our phylogenetic analyses of concatenated nucleotide and amino acid data sets provide
overwhelming support for the hypothesis that the Charales rather than the Coleochaetales
diverged just before the emergence of the land plants. Eight of the 23 proteins investigated
(Atp6, Cox1, MttB, Nad2, Nad4, Nad5, Nad6, and Rps4) individually support the monophyly of

Charales and land plants but cannot reject alternative hypotheses. Of these proteins, the one
encoded by the single mitochondrial gene analyzed by Karol et al. (2001), Nad5, provides the
highest bootstrap support (93%) for the sister-group relationships between the Charales and land
plants.

The dominant view has been that Charales, are the sister lineage, but an alternative hypothesis
supports the Zygnematales (often referred to as pond scum) as the sister lineage. From a
morphological standpoint, the relationship between charophytes and land plants tells a good
story: as the charophyte lineages diverge, their body plans grow increasingly complex from
unicellular (Mesostigmatales) to sarcinoid packets (Chlorokybales) to un-branched filaments
(Klebsormidiales) to branched filaments (Zygnematales), to parenchematous tissue
(Coleochaetales) and finally to the macrophytes (Charales). From there, the body plans evolve
into early land colonizers equipped with complex tissues allowing life out of water. Similarly,
sexual reproduction evolves from isogamy in the ancestral lineages to oogamy into the more
derived charophyte lineages. But in spite of morphological support for Charales as sister to land
plants, other data conflict with this interpretation.
Plastid gene phylogenies provide support for Zygnematales as sister to land plants. In addition,
new data based on nuclear genes support this alternative topology.
Sabina Wodniok et al in Origin of land plants: Do conjugating green
algae hold the key?
The researchers use a large data set of comprise of nuclear-encoded genes
(129 proteins) from 40 green plant taxa (Viridiplantae) including 21
embryophytes and six streptophyte algae, representing all major
streptophyte algal lineages.
Phylogenomic analyses of nuclear and chloroplast data indicate that the
Charales are most likely not the closest living extant relatives of the
embryophytes despite their morphological complexity. Instead, the analyses
favor either the Zygnematales or, less likely, a clade consisting of the
Zygnematales and Coleochaetales as the sister group of embryophytes.
It seems plausible that the simpler morphology of extant Zygnematales
represents a secondary simplification, similar to the loss of flagellate cells in
this group, which may actually represent an adaptation to ensure sexual
reproduction in the absence of free water. Alternatively, the morphological

complexity of the Charales and Coleochaetales might have evolved

independently after the three evolutionary lines (Coleochaetales,Charales,
and Zygnematales) diverged.
An extended taxon sampling and/ or analyses of larger data sets such as
complete genomes/ transcriptomes will likely be necessary to shed further
light on the intangible sister group of the embryophyte plants.

In line with this finding, another group of researchers come out with a more solid evidence
supporting Zygnematales as the closest living relatives to land plants. Ruth E. Timmel and some
other researchers were conducting a research to address the uncertainty of the sister lineages of
land plants. They sought a comprehensive genome scale analysis using a deep sampling of many
genes drawn from seven species distributed across all major charophyte lineages: Charales,
Coleochaetales, Zygnematales, Klebsormidiales, and Chlorokybales. The taxon sampling used
included a total of 14 taxa: eight charophytes, four land plants and two chlorophytes. This study,
which includes all charophyte lineages provides a robust, well-supported result that land plant
and Zygnematales are sister lineages. Previous hypotheses of increasing morphological
complexity
[20,21] are not congruent with the results of our study. However,
multiple gains and losses of multicellularity across all green algae
have been well documented, as has the reduction of characters in
the Zygnematales.
Evidence
1) Broad Phylogenomic Sampling and the Sister Lineage of Land Plants
Ruth E. Timme1, Tsvetan R. Bachvaroff, Charles F. Delwiche (a research article that
support the previous research)
The research paper provide a well supported, 160-nuclear-gene phylogenomic analysis
supporting the Zygnematales as the closest living relative to land plants.

Bryophytes as the first land plant

There is little doubt that land plants are monophyletic in origin, and that
bryophytes represent the first land plants. However, their exact phylogeny
remains unresolved, especially with regard to which group of bryophytes
(liverworts, mosses or hornworts) represents the earliest form of land plants.
Recent studies have also questioned the monophyly of liverworts
themselves.

Conclusion
Virtually all data support paraphyly of bryophytes. Genomic structural
information and at least some sequence data identify liverworts as the
earliest land plants. Relationships among mosses, hornworts and vascular
plant are not resolved at present. Monophyly of mosses and hornworts seem
to be well established, but not that of liverworts.
Transition from algae to bryophytes to pteridophytes

However, additional phylogenetic dilemmas are the evolution of bryophytes

from algae and the transition from these first land plants to the
pteridophytes.

Land plants (embryophytes) are most closely related to the Charophyceae, a

small group of predominantly freshwater green algae, within which either
Coleochaetales (,15 living species; Fig.1a) or Charales (,400 living
species;Fig. 1b), or a group containing both, is sister group to land plants.

Land-plant monophyly is supported by comparative morphology and gene

sequences (18S rRNA, mitochondrial DNA: cox III). Relationships among the
major basal living groups are uncertain but the best supported hypothesis
resolves liverworts (Fig. 1c) as basal and either mosses (Fig. 1e) or hornworts
(Fig. 1d) as the living sister group to vascular plants (tracheophytes). Less
parsimonious hypotheses recognize bryophyte monophyly and either a
sister-group relationship with vascular plants or an origin from within basal
vascular plants