Charophyta

2 Description

Charophyta is a division of freshwater green algae.[1]

The terrestrial plants, the Embryophyta emerged within
Charophyta, with the class Zygnematophyceae as a
sister group.[2][3][4] In some charophyte groups, such as
Zygnematophyceae or conjugating green algae, agellae
are absent and sexual reproduction does not involve freeswimming agellate sperm. Flagellate sperm, however,
are found in stoneworts (Charales) and Coleochaetales,
orders of parenchymatous charophytes that are the closest
relatives of the land plants, where agellate sperm are also
present in all except the conifers and owering plants.[5]
Fossil stoneworts of Devonian age that are similar to those
of the present day have been described from the Rhynie
chert of Scotland.[6]

The Zygnematophyceae or, as they used to be called,

Conjugatophyceae, generally possess two fairly elaborate
chloroplasts in each cell, rather than many discoid ones.
They reproduce asexually by the development of a septum between the two cell-halves or semi-cells (in unicellular forms, each daughter-cell develops the other semicell afresh) and sexually by conjugation, or the fusion of
the entire cell-contents of the two conjugating cells. The
saccoderm desmids and the placoderm or true desmids,
unicellular or lamentous members of the Zygnematophyceae, are dominant in non-calcareous, acid waters of
oligotrophic or primitive lakes (e.g. Wastwater), or in
lochans, tarns and bogs, as in the West of Scotland, Eire,
parts of Wales and of the Lake District.[10]

Klebsormidium, the type of the Klebsormidiophyceae, is

a simple lamentous form with circular, plate-like chloroplasts, reproducing by fragmentation, by dorsiventral, bi1 Classication
ciliate swarmers and, according to Wille, a twentiethcentury algologist, by aplanospores.[11] The same source
states that sexual reproduction is simple and isogamous
Charophyta are complex green algae that form a sis- (the gametes are outwardly identical without a distinct
ter group to the Chlorophyta and within which the male and female, at least to look at).
Embryophyta emerged. The chlorophyte and charophyte
green algae and the embryophytes or land plants form The various groups included in the Charophyta have dia clade called the green plants or Viridiplantae, that is verse and idiosyncratic reproductive systems, sometimes
united among other things by the absence of phycobilins, with complex reproductive organs. The unique habit
among the algae of protecting the overwintering zygote
the presence of chlorophyll b and chlorophyll a, cellulose in the cell wall and the use of starch, stored within the tissues of the parent gametophyte is one of
several characteristics of Coleochaetales that suggest that
in the plastids, as a storage polysaccharide. Unlike
[12]
chlorophytes, the charophytes and embryophytes share they are a sister group to the embryophytes.
several traits, such as the presence of certain enzymes The Charales or stoneworts are freshwater algae with
(class I aldolase, Cu/Zn Superoxide dismutase, Glycolate slender green or grey stems; the grey colour of many
Oxidase, agellar peroxidase), lateral agella (when species results from the deposition of lime on the walls,
present), and, in many species, the use of phragmoplasts masking the green colour of the chlorophyll. The main
in mitosis. [7] Thus Charophyta and Embryophyta stems are slender and branch occasionally. Lateral
together form the clade Streptophyta, excluding the branchlets occur in whorls at regular intervals up the stem,
Chlorophyta. The Charophyta sensu lato does not they are attached by rhizoids to the substrate.[13] The
include all the descendents of their common ances- reproductive organs consist of antheridia and oogonia,
tor with the embryophytes and as such they form a though the structures of these organs dier considerably
paraphyletic group i.e. the classes Chlorokybophyceae; from the corresponding organs in other algae. As a reMesostigmatophyceae;
Klebsormidiophyceae; sult of fertilization a protonema is formed, from which
Zygnematophyceae;
Coleochaetophyceae
and the sexually reproducing algae develops.
Chaetosphaeridiales are removed from the division.[8]
Charophytes are frequently found in hard water with disCharophytes such as Palaeonitella cranii and possibly the
yet unassigned Parka decipiens[9] are present in the fossil record of the Devonian.[6] Palaeonitella diered little
from some present-day stoneworts.

solved calcium or magnesium carbonates. They tolerate low concentrations of salt, and are found in the inner reaches of the Baltic Sea[14] and in tropical brackish lagoons[15] but not in marine environments. The wa1

REFERENCES

ter must be still, or only slow-owing, oligotrophic or development of the antheridium before the formation of
mesotrophic and little pollution due to sewage.
the oogonium, thus preventing fertilization by antheroThe Charophyceae are obligate aquatic algae, growing zoids of the same alga. In this case the two types of sex
submerged in calcareous fresh water. They are dis- organs usually arise from dierent points on the lateral
tributed throughout the world from the tropics to cold branches
temperate zones.
Six genera are recognized:Chara
Lamprothamnium
Lychnothamnus
Nitella
Nitellopsis
Tolypella.

2.1

[13]

Cell structure

There are numerous small discoid chloroplasts, which are

disposed around the periphery of the cells. No pyrenoids
are present. The large internodal cells are sometimes
multinucleate, and their nuclei often possess large nucleoli and scanty chromatin. In these cells the cytoplasm
forms only a peripheral layer with a large central vacuole. The cell walls are composed of cellulose, though
there may be also a supercial layer of a more gelatinous
material of unknown composition.
The storage material is starch, except in the oospore,
where oil also occurs. This starch also accumulates in
special storage structures, termed bulbils, which consist
of rounded cells of varying size which are developed in
clusters on the lower stem and root nodes. They are
mainly developed when the algae are growing in ne slimy
mud.
Cytoplasmic streaming was rst demonstrated in the giant
cells of Chara internodes by Giovanni Battista Amici, in
1818.

2.2

Sexual reproduction

The reproductive organs of the Charales show a high degree of specialization. The female organ is a large oval
structure with an envelope of spirally arranged, bright
green laments of cells. It is termed an oogonium. The
male organ is also large, bright yellow or red in colour,
spherical in shape, and is usually termed an antheridium,
though some workers regard it as a multiple structure
rather than a single organ. The sex organs are developed
in pairs from the adaxial nodal cell at the upper nodes of
the primary lateral branches, the oogonium being formed
above the antheridium. They are suciently large to be
easily seen with the naked eye, especially the bright orange or red antheridium. Many species are dioecious. In
others the monoecious condition is complicated by the

2.3 Vegetative propagation

Vegetative propagation occurs readily in the Charales.
Secondary protonemata may develop even more rapidly
than primary ones. Fragments of nodes, dormant cells
of algae after hibernation or the basal nodes of primary
rhizoids may all produce these secondary protonemata,
from which fresh sexual algae can arise. It is probably
this power of vegetative propagation which explains the
fact that species of Characeae are generally found forming dense clonal mats in the beds of ponds or streams,
covering quite large areas.

3 References
[1] Lewis, Louise A.; Richard M. McCourt (2004).
Green algae and the origin of land plants (abstract).
American Journal of Botany. 91 (10): 15351556.
doi:10.3732/ajb.91.10.1535. PMID 21652308.
[2] BMC Evolutionary Biology 2014, 14:23.
[3] Wickett, Norman J.; Mirarab, Siavash; Nguyen, Nam;
Warnow, Tandy; Carpenter, Eric; Matasci, Naim;
Ayyampalayam, Saravanaraj; Barker, Michael S.;
Burleigh, J. Gordon (2014-11-11). Phylotranscriptomic
analysis of the origin and early diversication
of land plants.
Proceedings of the National
Academy of Sciences.
111 (45): E4859E4868.
doi:10.1073/pnas.1323926111. ISSN 0027-8424. PMC
4234587 . PMID 25355905.
[4] Vries, Jan de; Stanton, Amanda; Archibald, John M.;
Gould, Sven B. (2016-02-16). Streptophyte Terrestrialization in Light of Plastid Evolution. Trends in Plant
Science. 0 (0). doi:10.1016/j.tplants.2016.01.021. ISSN
1360-1385.
[5] Vaughn, K.C.; Renzaglia, K.S. (2006). Structural
and immunocytochemical characterization of the Ginkgo
biloba L. sperm motility apparatus.. Protoplasma. 227
(2-4): 16573. doi:10.1007/s00709-005-0141-3.
[6] Kelman, R.; Feist, M.; Trewin, N.H.; Hass, H. (2003).
Charophyte algae from the Rhynie chert. Transactions
of the Royal Society of Edinburgh: Earth Sciences. 94 (4):
445455. doi:10.1017/s0263593300000808.
[7] Leliaert, F., Smith, D.R., Moreau, H., Herron, M.D., Verbruggen, H., Delwiche, C.F. & De Clerck, O. (2012).
Phylogeny and molecular evolution of the green algae
(PDF). Critical Reviews in Plant Sciences. 31: 146.
doi:10.1080/07352689.2011.615705.
[8] Tree of Life: Green plants

[9] Hemsley, A.R. (1989). The ultrastructure of the spores

of the Devonian plant Parka decipiens". Annals of Botany.
64 (3): 359367.
[10] West, Fritsch, G.S., F.E. (1927). A Treatise of the British
Freshwater Algae. Cambridge: Cambridge University
Press.
[11] Fritsch, F.E. (1935). The Structure and Reproduction of
the Algae, vol I. Cambridge University Press. pp. 205
206.
[12] Becker, B.; Marin, B. (2009). Streptophyte algae and
the origin of embryophytes. Annals of Botany. 103 (7):
9991004. doi:10.1093/aob/mcp044. PMC 2707909 .
PMID 19273476.
[13] Bryant 2007, J. The Stoneworts (Chlorophyta, Charales)
in Guiry, M.D., John, D.M., Rindi, F. and McCarthy, T.K
(Ed) New Survey of Clare Island Volume 6: The Freshwater and Terrestrial Algae. Royal Irish Academy. ISBN
9781904890317
[14] Schubert, H.; Blindow, I. (2004). Charophytes of the
Baltic Sea. Gantner Verlag. ISBN 3906166066.
[15] Palma-Silva, C.; Albertoni, E.F.; Esteves, F.A. (2004).
Charophytes as nutrient and energy reservoir in a tropical
coastal lagoon impacted by humans (RJ, Brazil).. Brazilian Journal of Biology. 64: 479487. doi:10.1590/s151969842004000300011.

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