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''Chara'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
charophyte Charophyta () is a paraphyletic group of freshwater green algae, called charophytes (), sometimes treated as a division, yet also as a superdivision. The terrestrial plants, the Embryophyta emerged deep within Charophyta, possibly from terres ...
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
in the family
Characeae Characeae is a family of freshwater green algae in the order Charales, commonly known as stoneworts. They are also known as brittleworts or skunkweed, from the fragility of their lime-encrusted stems, and from the foul odor these produce when ste ...
. They are multicellular and superficially resemble land plants because of
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
-like and
leaf A leaf (: leaves) is a principal appendage of the plant stem, stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leav ...
-like structures. They are found in freshwater, particularly in limestone areas throughout the northern temperate zone, where they grow submerged, attached to the muddy bottom. They prefer less oxygenated and hard water and are not found in waters where mosquito larvae are present. They are covered with
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
(CaCO3) deposits and are commonly known as stoneworts.
Cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
have been found growing as epiphytes on the surfaces of ''Chara'', where they may be involved in fixing
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
, which is important to plant nutrition.


Structure

The branching system of ''Chara'' species is complex with branches derived from apical cells which cut off segments at the base to form nodal and internodal cells alternately.Round, F.E. 1965.''The Biology of the algae.'' Ernest Arnold. The main axes bear whorls of branches in a superficial resemblance to ''
Equisetum ''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which ...
'' (a vascular plant). They are typically anchored to the
littoral The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely i ...
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
by means of branching underground
rhizoid Rhizoids are protuberances that extend from the lower epidermal cells of bryophytes and algae. They are similar in structure and function to the root hairs of vascular land plants. Similar structures are formed by some fungi. Rhizoids may be un ...
s. ''Chara'' plants are rough to the touch because of deposited
calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
salts on the cell wall. The metabolic processes associated with this deposition often give ''Chara ''plants a distinctive and unpleasant smell of
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is toxic, corrosive, and flammable. Trace amounts in ambient atmosphere have a characteristic foul odor of rotten eggs. Swedish chemist ...
.


Morphology

The plant body is a
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
. It consists of the main axis (differentiated into nodes and internodes), dimorphic branches (long branch of unlimited growth and short branches of limited growth), rhizoids (multicellular with oblique septa) and stipulodes (needle-shaped structures at the base of secondary laterals).


Reproduction

''Chara'' reproduces vegetatively and sexually. Vegetative reproduction takes place by
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
s, amylum stars and secondary
protonema A protonema (plural: protonemata) is a thread-like chain of cells that forms the earliest stage of development of the gametophyte (the haploid phase) in the life cycle of mosses. When a moss first grows from a spore, it starts as a ''germ tube'', ...
ta. The sex organs are a multicellular and jacketed globule or antheridium (male) and nucule or archegonium (female). The antheridia and archegonia may occur on separate plants (
dioicy Dioicy () is a sexual system in non-vascular plants where archegonia (female organs) and antheridia (male organs) are produced on separate plants in the gametophyte phase. It is one of the two main sexual systems in bryophytes, the other bei ...
), together on the same plant (conjoined monoicy) or separately on the same plant (sejoined monoicy). The reproductive organs of the Charales show a high degree of specialization. The female organ, called an
oogonium An oogonium (: oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in large ...
is a large oval structure with an envelope of spirally arranged, bright green filaments of cells. The male organ or is also large, bright yellow or red in colour, spherical in shape, and is usually termed an
antheridium An antheridium is a haploid structure or organ producing and containing male gametes (called ''antherozoids'' or sperm). The plural form is antheridia, and a structure containing one or more antheridia is called an androecium. The androecium is al ...
, 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 sufficiently 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 development of the antheridium before the formation of the oogonium, thus preventing fertilization by antherozoids of the same alga. In this case the two types of sex organs usually arise from different points on the lateral branches. All cells of the Charales are
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
except for the fertilized
zygote A zygote (; , ) is a eukaryote, eukaryotic cell (biology), cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individ ...
, the large single cell in the interior of the oogonium, which becomes enclosed in a thickened hard wall to form an
oospore An oospore is a thick-walled sexual spore that develops from a fertilized oosphere in some algae, fungi, and oomycetes. They are believed to have evolved either through the fusion of two species or the chemically induced stimulation of mycelia ...
that awaits favorable conditions for germination. Upon germination the diploid oospore undergoes
meiosis Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
, producing four haploid nuclei. A septum divides a small apical cell with one haploid nucleus from a large basal cell containing the other three nuclei, which will slowly degenerate. The oospore apical cell divides to produce the protonemal initial, from which the primary protonema arises, and the rhizoidal initial, from which the primary rhizoid descends. From these the alga continues its development.


Species


Distribution

''Chara'' has a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
, from 69 degrees north in northern Norway to about 49 degrees south in
Kerguelen Islands The Kerguelen Islands ( or ; in French commonly ' but officially ', ), also known as the Desolation Islands (' in French), are a group of islands in the subantarctic, sub-Antarctic region. They are among the Extremes on Earth#Remoteness, most i ...
(Pal et al., 1962). About 27 species are found in
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
.Soni, N.K. Fundamentals of Botany Vol. 1, Tata McGraw-Hill Education, p72, There are about 40 species of Chara in Europe, where they are commonly found in the specific
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
-type designated as H3140 (hard oligo-mesotrophic waters with benthic vegetation of ''Chara spp'' h1) in the
Natura 2000 Natura 2000 is a network of nature protection areas in the territory of the European Union. It is made up of Special Areas of Conservation and Special Protection Areas designated under the Habitats Directive and the Birds Directive, respectiv ...
plans of the European Union. Although this habitat is found all across Europe, it is threatened and to be protected and preserved.
The Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
are home to 20 species of ''Chara'', growing in lakes and ponds of the habitat-type H3140. The H3140 habitats in the Netherlands, are considered important in the overall preservation efforts and therefore also for the ''Chara'' species in general.
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
. Here, many former ''Chara'' habitats (H3140) have been polluted by either toxins or excessive amounts of nutrients (in particular
phosphate Phosphates are the naturally occurring form of the element phosphorus. In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthop ...
s and
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
), but a few large lakes and ponds remain. ''Chara'' is found growing in the very clean
hard water Hard water is water that has a high mineral content (in contrast with "soft water"). Hard water is formed when water percolates through deposits of limestone, chalk or gypsum, which are largely made up of calcium and magnesium carbonates, bic ...
lakes of
Thy National Park Thy National Park () is a national park area in Thy, Denmark, opened to the public on 22 August 2008. It is located in Northwest Jutland, along the coast from Hanstholm to Agger Tange and it spans north to south and east to west. The total are ...
like Nors Sø for example.
Tissø Tissø is the 4th largest freshwater lake in Denmark, at 12.3 km2. It is located on the western part of Zealand, in the municipality of Kalundborg municipality, Kalundborg. There are several small towns and villages near the lake, of which ...
lake (fourth-largest lake in Denmark) is also a H3140 habitat and contains ''Chara'' species. Britain & Ireland: - C. aspera, C. baltica, C. braunii, C. canescens, C. connivens, C. fragifera, C. globularis, C. hispida, C. muscosa, C. pedunculata, C. tomenosa, C. vulgaris (as at 1986)
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
:- Co. Galway. Eglinton Canal '' Chara virgata'' Kütz., '' Chara rudis'' (A.Braun) Leonhardii and ''
Nitella flexilis ''Nitella flexilis'', the smooth stonewort, is a freshwater species of characean algae that is used as a model organism for its large cell size and relative ease of cultivation in the laboratory. Description A robust species growing up to a me ...
'' (L.) C.Agardh.Pybus, C. and O'Halloran, P. 2009. Distribution of some submerged aquatic macrophytes in Eglinton Canal, Galway. ''Ir. Nat. J.'' 30: 51 – 53


References

{{Authority control Charophyta Charophyta genera Extant Silurian first appearances