Charophyta

Charophyta are complex green algae that form a sister group to the Chlorophyta and within which the Embryophyta emerged. The chlorophyte and charophyte green algae and the embryophytes or land plants form a clade called the green plants or Viridiplantae, that is united among other things by the absence of phycobilins, the presence of chlorophyll a and chlorophyll b, cellulose in the cell wall and the use of starch, stored in the plastids, as a storage polysaccharide. The charophytes and embryophytes share several traits that distinguish them from the chlorophytes, such as the presence of certain enzymes (class I aldolase, Cu/Zn superoxide dismutase, glycolate oxidase, flagellar peroxidase), lateral flagella (when present), and, in many species, the use of phragmoplasts in mitosis.[13] Thus Charophyta and Embryophyta together form the clade Streptophyta, excluding the Chlorophyta.

Charophytes such as Palaeonitella cranii and possibly the yet unassigned Parka decipiens[14] are present in the fossil record of the Devonian.[12] Palaeonitella differed little from some present-day stoneworts.

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 semi-cell 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 filamentous 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.[26]

Klebsormidium, the type of the Klebsormidiophyceae, is a simple filamentous form with circular, plate-like chloroplasts, reproducing by fragmentation, by dorsiventral, biciliate swarmers and, according to Wille, a twentieth-century algologist, by aplanospores.[27] Sexual reproduction is simple and isogamous (the male and female gametes are outwardly indistinguishable).[27]

The various groups included in the Charophyta have diverse and idiosyncratic reproductive systems, sometimes with complex reproductive organs. The unique habit among the algae of protecting the overwintering zygote within the tissues of the parent gametophyte is one of several characteristics of Coleochaetales that suggest that they are a sister group to the embryophytes.[28]

The Charales or stoneworts are freshwater algae with slender green or grey stems; the grey colour of many species results from the deposition of lime on the walls, masking the green colour of the chlorophyll. The main stems are slender and branch occasionally. Lateral branchlets occur in whorls at regular intervals up the stem, they are attached by rhizoids to the substrate.[29] The reproductive organs consist of antheridia and oogonia, though the structures of these organs differ considerably from the corresponding organs in other algae. As a result of fertilization a protonema is formed, from which the sexually reproducing algae develops.

Charophytes are frequently found in hard water with dissolved calcium or magnesium carbonates. They tolerate low concentrations of salt, and are found in the inner reaches of the Baltic Sea[30] and in tropical brackish lagoons[31] but not in marine environments. The water must be still, or only slow-flowing, oligotrophic or mesotrophic and little pollution due to sewage.

The Charophyceae are obligate aquatic algae, growing submerged in calcareous fresh water. They are distributed throughout the world from the tropics to cold temperate zones.

Six genera are recognized:

• Chara
• Lamprothamnium
• Lychnothamnus
• Nitella
• Nitellopsis
• Tolypella.[29]

Sexual reproduction

Light micrograph of a whole-mount slide of an oogonium (or nucule) and antheridium (or globule) of Chara (conjoined monoecy)

The reproductive organs of the Charales show a high degree of specialization. The female organ, called an oogonium is a large oval structure with an envelope of spirally arranged, bright green filaments of cells. It is termed an oogonium. The male organ or 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 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 except for the fertilized zygote, the large single cell in the interior of the oogonium, which becomes enclosed in a thickened hard wall to form an oospore that awaits favorable conditions for germination. Upon germination the diploid oospore undergoes meiosis, 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.[32]

• Data related to Charophyta at Wikispecies