Green algae

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Green algae
Green algae.png
Green algal diversity. From top left corner: Picocystis (Picocystophyceae), Acetabularia (Ulvophyceae), Botryococcus (Trebouxiophyceae), Volvox (Chlorophyceae), Klebsormidium (Klebsormidiophyceae), Chara (Charophyceae), Spirogyra and Micrasterias (Zygnematophyceae)
Scientific classification OOjs UI icon edit-ltr.svg
Clade: Archaeplastida
Clade: Viridiplantae
Groups included
Cladistically included but traditionally excluded taxa

The green algae (sg.: 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 (Embryophytes) have emerged deep in the Charophyte alga as a sister of the Zygnematophyceae. [1] [2] [3] Since the realization that the Embryophytes emerged within the green algae, some authors are starting to include them. [2] [4] [5] [6] [7] [ excessive citations ] The completed clade that includes both green algae and embryophytes is monophyletic and is referred to as the clade Viridiplantae and as the kingdom Plantae. The green algae include unicellular and colonial flagellates, most with two flagella per cell, as well as various colonial, coccoid (spherical), and filamentous forms, and macroscopic, multicellular seaweeds. There are about 22,000 species of green algae, [8] many of which live most of their lives as single cells, while other species form coenobia (colonies), long filaments, or highly differentiated macroscopic seaweeds.

Contents

A few other organisms rely on green algae to conduct photosynthesis for them. The chloroplasts in dinoflagellates of the genus Lepidodinium , euglenids and chlorarachniophytes were acquired from ingested endosymbiont green algae, [9] and in the latter retain a nucleomorph (vestigial nucleus). Green algae are also found symbiotically in the ciliate Paramecium , and in Hydra viridissima and in flatworms. Some species of green algae, particularly of genera Trebouxia of the class Trebouxiophyceae and Trentepohlia (class Ulvophyceae), can be found in symbiotic associations with fungi to form lichens. In general the fungal species that partner in lichens cannot live on their own, while the algal species is often found living in nature without the fungus. Trentepohlia is a filamentous green alga that can live independently on humid soil, rocks or tree bark or form the photosymbiont in lichens of the family Graphidaceae. Also the macroalga Prasiola calophylla (Trebouxiophyceae) is terrestrial, [10] and Prasiola crispa , which live in the supralittoral zone, is terrestrial and can in the Antarctic form large carpets on humid soil, especially near bird colonies. [11]

Cellular structure

Green algae have chloroplasts that contain chlorophyll a and b, giving them a bright green colour, as well as the accessory pigments beta carotene (red-orange) and xanthophylls (yellow) in stacked thylakoids. [12] [13] The cell walls of green algae usually contain cellulose, and they store carbohydrate in the form of starch. [14]

All green algae have mitochondria with flat cristae. When present, paired flagella are used to move the cell. They are anchored by a cross-shaped system of microtubules and fibrous strands. Flagella are only present in the motile male gametes of charophytes [15] bryophytes, pteridophytes, cycads and Ginkgo , but are absent from the gametes of Pinophyta and flowering plants.

Members of the class Chlorophyceae undergo closed mitosis in the most common form of cell division among the green algae, which occurs via a phycoplast. [16] By contrast, charophyte green algae and land plants (embryophytes) undergo open mitosis without centrioles. Instead, a 'raft' of microtubules, the phragmoplast, is formed from the mitotic spindle and cell division involves the use of this phragmoplast in the production of a cell plate. [17]

Origins

Photosynthetic eukaryotes originated following a primary endosymbiotic event, where a heterotrophic eukaryotic cell engulfed a photosynthetic cyanobacterium-like prokaryote that became stably integrated and eventually evolved into a membrane-bound organelle: the plastid. [18] This primary endosymbiosis event gave rise to three autotrophic clades with primary plastids: the (green) plants (with chloroplasts) the red algae (with rhodoplasts) and the glaucophytes (with muroplasts). [19]

Evolution and classification

A growth of the green seaweed Ulva on rock substratum at the ocean shore; some green seaweeds like Ulva are quick to utilize inorganic nutrients from land runoff, and thus can be indicators of nutrient pollution Intertidal greenalgae.jpg
A growth of the green seaweed Ulva on rock substratum at the ocean shore; some green seaweeds like Ulva are quick to utilize inorganic nutrients from land runoff, and thus can be indicators of nutrient pollution

Green algae are often classified with their embryophyte descendants in the green plant clade Viridiplantae (or Chlorobionta). Viridiplantae, together with red algae and glaucophyte algae, form the supergroup Primoplantae, also known as Archaeplastida or Plantae sensu lato. The ancestral green alga was a unicellular flagellate. [20]

The Viridiplantae diverged into two clades. The Chlorophyta include the early diverging prasinophyte lineages and the core Chlorophyta, which contain the majority of described species of green algae. The Streptophyta include charophytes and land plants. Below is a consensus reconstruction of green algal relationships, mainly based on molecular data. [21] [20] [22] [23] [5] [24] [25] [26] [27] [28] [29] [30] [1] [ excessive citations ]

Viridiplantae/
green algae

The basal character of the Mesostigmatophyceae, Chlorokybophyceae and spirotaenia are only more conventionally basal Streptophytes.

The algae of this paraphyletic group "Charophyta" were previously included in Chlorophyta, so green algae and Chlorophyta in this definition were synonyms. As the green algae clades get further resolved, the embryophytes, which are a deep charophyte branch, are included in "algae", "green algae" and "Charophytes", or these terms are replaced by cladistic terminology such as Archaeplastida, Plantae/Viridiplantae, and streptophytes, respectively. [31]

Reproduction

Green algae conjugating Algaeconjugate.jpg
Green algae conjugating

Green algae are a group of photosynthetic, eukaryotic organisms that include species with haplobiontic and diplobiontic life cycles. The diplobiontic species, such as Ulva , follow a reproductive cycle called alternation of generations in which two multicellular forms, haploid and diploid, alternate, and these may or may not be isomorphic (having the same morphology). In haplobiontic species only the haploid generation, the gametophyte is multicellular. The fertilized egg cell, the diploid zygote, undergoes meiosis, giving rise to haploid cells which will become new gametophytes. The diplobiontic forms, which evolved from haplobiontic ancestors, have both a multicellular haploid generation and a multicellular diploid generation. Here the zygote divides repeatedly by mitosis and grows into a multicellular diploid sporophyte. The sporophyte produces haploid spores by meiosis that germinate to produce a multicellular gametophyte. All land plants have a diplobiontic common ancestor, and diplobiontic forms have also evolved independently within Ulvophyceae more than once (as has also occurred in the red and brown algae). [32]

Diplobiontic green algae include isomorphic and heteromorphic forms. In isomorphic algae, the morphology is identical in the haploid and diploid generations. In heteromorphic algae, the morphology and size are different in the gametophyte and sporophyte. [33]

Reproduction varies from fusion of identical cells (isogamy) to fertilization of a large non-motile cell by a smaller motile one (oogamy). However, these traits show some variation, most notably among the basal green algae called prasinophytes.

Haploid algal cells (containing only one copy of their DNA) can fuse with other haploid cells to form diploid zygotes. When filamentous algae do this, they form bridges between cells, and leave empty cell walls behind that can be easily distinguished under the light microscope. This process is called conjugation and occurs for example in Spirogyra .

Sex pheromone

Sex pheromone production is likely a common feature of green algae, although only studied in detail in a few model organisms. Volvox is a genus of chlorophytes. Different species form spherical colonies of up to 50,000 cells. One well-studied species, Volvox carteri (2,000 – 6,000 cells) occupies temporary pools of water that tend to dry out in the heat of late summer. As their environment dries out, asexual V. carteri quickly die. However, they are able to escape death by switching, shortly before drying is complete, to the sexual phase of their life cycle that leads to production of dormant desiccation-resistant zygotes. Sexual development is initiated by a glycoprotein pheromone (Hallmann et al., 1998). This pheromone is one of the most potent known biological effector molecules. It can trigger sexual development at concentrations as low as 10−16M. [34] Kirk and Kirk [35] showed that sex-inducing pheromone production can be triggered experimentally in somatic cells by heat shock. Thus heat shock may be a condition that ordinarily triggers sex-inducing pheromone in nature. [34]

The Closterium peracerosum-strigosum-littorale (C. psl) complex is a unicellular, isogamous charophycean alga group that is the closest unicellular relative to land plants. Heterothallic strains of different mating type can conjugate to form zygospores. Sex pheromones termed protoplast-release inducing proteins (glycopolypeptides) produced by mating-type (-) and mating-type (+) cells facilitate this process. [36]

Physiology

The green algae, including the characean algae, have served as model experimental organisms to understand the mechanisms of the ionic and water permeability of membranes, osmoregulation, turgor regulation, salt tolerance, cytoplasmic streaming, and the generation of action potentials. [37]

Related Research Articles

<span class="mw-page-title-main">Chlorophyta</span> Phylum of green algae

Chlorophyta is a division of green algae informally called chlorophytes.

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

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 sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

<i>Volvox</i> Genus of algae

Volvox is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. Volvox species form spherical colonies of up to 50,000 cells, and for this reason they are sometimes called globe algae. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 1700. Volvox diverged from unicellular ancestors approximately 200 million years ago.

<span class="mw-page-title-main">Alternation of generations</span> Reproductive cycle of plants and algae

Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.

<span class="mw-page-title-main">Brown algae</span> Large group of multicellular algae, comprising the class Phaeophyceae

Brown algae are a large group of multicellular algae comprising the class Phaeophyceae. They include many seaweeds located in colder waters of the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and polar regions. Many brown algae, such as members of the order Fucales, commonly grow along rocky seashores. Most brown algae live in marine environments, where they play an important role both as food and as a potential habitat. For instance, Macrocystis, a kelp of the order Laminariales, may reach 60 m (200 ft) in length and forms prominent underwater kelp forests that contain a high level of biodiversity. Another example is Sargassum, which creates unique floating mats of seaweed in the tropical waters of the Sargasso Sea that serve as the habitats for many species. Some members of the class, such as kelps, are used by humans as food.

<span class="mw-page-title-main">Bryophyte</span> Terrestrial plants that lack vascular tissue

Bryophytes are a group of land plants (embryophytes), sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants: the liverworts, hornworts, and mosses. In the strict sense, the division Bryophyta consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although some species can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce sexually by spores and asexually by fragmentation or the production of gemmae.

<span class="mw-page-title-main">Embryophyte</span> Subclade of green plants, also known as land plants

The embryophytes are a clade of plants, also known as Embryophyta or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of freshwater charophyte green algae as a sister taxon of Charophyceae, Coleochaetophyceae and Zygnematophyceae. Embryophytes consist of the bryophytes and the polysporangiophytes. Living embryophytes include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and angiosperms. Embryophytes have diplobiontic life cycles.

<span class="mw-page-title-main">Biological life cycle</span> Series of stages of an organism

In biology, a biological life cycle is a series of stages of the life of an organism, that begins as a zygote, often in an egg, and concludes as an adult that reproduces, producing an offspring in the form of a new zygote which then itself goes through the same series of stages, the process repeating in a cyclic fashion.

<span class="mw-page-title-main">Sporophyte</span> Diploid multicellular stage in the life cycle of a plant or alga

A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.

<span class="mw-page-title-main">Streptophyta</span> Clade consisting of the charophyte algae and land plants

Streptophyta, informally the streptophytes, is a clade of plants. The composition of the clade varies considerably between authors, but the definition employed here includes land plants and all green algae except the Chlorophyta and the more basal Prasinodermophyta.

<span class="mw-page-title-main">Charophyta</span> Phylum of algae

Charophyta is a group of freshwater green algae, called charophytes, sometimes treated as a division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged deep within Charophyta, possibly from terrestrial unicellular charophytes, with the class Zygnematophyceae as a sister group.

<i>Cladophora</i> Genus of filamentous green algae

Cladophora is a genus of reticulated filamentous green algae in the class Ulvophyceae. They may be referred to as reticulated algae, branching algae, or blanket weed. The genus has a worldwide distribution and is harvested for use as a food and medicine.

<span class="mw-page-title-main">Viridiplantae</span> Clade of archaeplastids including green algae and the land plants

Viridiplantae constitute a clade of eukaryotic organisms that comprises approximately 450,000–500,000 species that play important roles in both terrestrial and aquatic ecosystems. They include the green algae, which are primarily aquatic, and the land plants (embryophytes), which emerged from within them. Green algae traditionally excludes the land plants, rendering them a paraphyletic group. However it is accurate to think of land plants as a kind of alga. Since the realization that the embryophytes emerged from within the green algae, some authors are starting to include them. They have cells with cellulose in their cell walls, and primary chloroplasts derived from endosymbiosis with cyanobacteria that contain chlorophylls a and b and lack phycobilins. Corroborating this, a basal phagotroph archaeplastida group has been found in the Rhodelphydia.

<span class="mw-page-title-main">Archaeplastida</span> Clade of eukaryotes containing land plants and some algae

The Archaeplastida are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans. The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by phagocytosis of a cyanobacterium. All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae. Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.

The Mesostigmatophyceae are a class of basal green algae found in freshwater. In a narrow circumscription, the class contains a single genus, Mesostigma. AlgaeBase then places the order within its circumscription of Charophyta. A clade containing Chlorokybus and Spirotaenia may either be added, or treated as a sister, with Chlorokybus placed in a separate class, Chlorokybophyceae. When broadly circumscribed, Mesostigmatophyceae may be placed as sister to all other green algae, or as sister to all Streptophyta.

Mesostigma is a genus of unicellular biflagellate freshwater green algae, with a single species Mesostigma viride, covered by an outer layer of basket‐like scales instead of a cell wall. It is the only known genus in the class Mesostigmatophyceae.

<i>Coleochaete</i> Genus of algae

Coleochaete is a genus of parenchymatous charophyte green algae in the order Coleochaetales. They are haploid, reproduce both sexually and asexually, and have true multicellular organisation, with plasmodesmata communicating between adjacent cells. The plants form flat, sprawling discs on solid surfaces in freshwater streams worldwide, usually as epiphytes on aquatic plants or growing on the surface of stones. They are seen as one of two most probable sister groups to land plant species, the second candidate group being the Characeae. The issue is still not resolved. As they show some of the earliest and simplest features of multicellular plant growth, they are ideal model organisms in the field of synthetic biology. They are easy to culture and techniques that have been used to study Arabidopsis thaliana are now being applied to Coleochaete.

<span class="mw-page-title-main">Klebsormidiaceae</span> Family of algae

The Klebsormidiaceae are a family containing five genera of charophyte green alga forming multicellular, non-branching filaments. The genus Chlorokybus was previously included as well, but this problematic and poorly known genus is now placed in a separate class Chlorokybophyceae.

<span class="mw-page-title-main">Red algae</span> Division of plant life

Red algae, or Rhodophyta, make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest phyla of algae, containing over 7,000 recognized species within over 900 genera amidst ongoing taxonomic revisions. The majority of species (6,793) are Florideophyceae, and mostly consist of multicellular, marine algae, including many notable seaweeds. Red algae are abundant in marine habitats. Approximately 5% of red algae species occur in freshwater environments, with greater concentrations in warmer areas. Except for two coastal cave dwelling species in the asexual class Cyanidiophyceae, no terrestrial species exist, which may be due to an evolutionary bottleneck in which the last common ancestor lost about 25% of its core genes and much of its evolutionary plasticity.

<span class="mw-page-title-main">Phragmoplastophyta</span> Clade of algae

The Phragmoplastophyta are a proposed sister clade of the Klebsormidiaceae in the Streptophyte/Charophyte clade. The Phragmoplastophyta consist of the Charophyceae and another unnamed clade which contains the Coleochaetophyceae, Zygnematophyceae, Mesotaeniaceae, and Embryophytes. It is an important step in the emergence of land plants within the green algae. It is equivalent to the ZCC clade/grade, cladistically granting the Embryophyta.

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