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Green algae

Green algae form a diverse group of primarily aquatic photosynthetic eukaryotes, including the division Chlorophyta and charophyte algae within Streptophyta, characterized by the presence of chlorophylls a and b, β-carotene, and other xanthophylls, which enable oxygenic photosynthesis akin to that in embryophytes (land plants).^[1] They store energy as starch within plastids and exhibit a wide array of morphologies, ranging from unicellular motile cells and non-motile coccoids to colonial, filamentous, and parenchymatous thalli that can form macroscopic seaweeds.^[2] Comprising approximately 22,000 described species, green algae inhabit marine, freshwater, and terrestrial environments, including extreme settings like hot deserts, polar regions, and symbiotic associations in lichens or within other organisms.^[3] Ecologically significant as primary producers, they contribute substantially to global oxygen production and serve as foundational components in aquatic food webs, while some species have applications in biotechnology for biofuels, pharmaceuticals, and food.^[4] Phylogenetically, green algae belong to the core green plants (Viridiplantae), with Chlorophyta forming a monophyletic clade and streptophytes (which include charophyte algae and land plants) the sister group, their lineages diverging approximately 1.5 billion years ago.^[1] The Chlorophyta division is subdivided into several classes based on molecular and ultrastructural data, including the flagellate Prasinophyceae (often considered ancestral), the marine and freshwater Ulvophyceae (encompassing siphonous and blade-like forms), the soil- and lichen-dwelling Trebouxiophyceae, and the highly diverse Chlorophyceae (featuring orders like Chlamydomonadales and Sphaeropleales).^[2] This classification highlights their evolutionary radiation, with cell walls varying from cellulose-based structures in advanced groups to glycoprotein scales in primitive ones, reflecting adaptations to diverse habitats.^[1] Green algae are pivotal in understanding plant evolution, as streptophyte lineages such as the Charophyceae (e.g., Chara and Coleochaete) share complex traits with land plants, including phragmoplast-mediated cell division, pectin-rich cell walls, and phytochrome photoreceptors, underscoring their role as ancestral to terrestrial flora.^[1] Their biodiversity hotspots occur in tropical marine realms and temperate freshwater systems, though cryptic species diversity revealed by genomics suggests higher richness than currently documented.^[5] Notable examples include the bloom-forming Ulva (sea lettuce) in coastal waters and the invasive Caulerpa taxifolia in Mediterranean seas, illustrating both ecological benefits and challenges posed by this group.^[4]

Introduction and Overview

Definition and Diversity

Green algae are photosynthetic eukaryotes characterized by the presence of chlorophylls a and b, which impart their characteristic green color, and are members of the Viridiplantae clade, a major lineage of plants that also encompasses land plants (embryophytes) but excludes them from the algal group.^[6] These organisms possess chloroplasts derived from a primary endosymbiosis event in which an ancestral eukaryote engulfed a cyanobacterium, and they typically feature cellulose in their cell walls, distinguishing them from other algal groups.^[6] Unlike land plants, green algae lack the complex adaptations for terrestrial life, such as extensive vascular systems, and are predominantly aquatic or subaerial in habitat. The diversity of green algae is remarkable, encompassing approximately 6,900 described species (as of 2024) that span a broad spectrum of morphological forms, from simple unicellular flagellates to highly organized multicellular structures.^[7] Unicellular representatives include species like Chlamydomonas reinhardtii, a motile freshwater alga often used in laboratory studies.^[6] Colonial forms, such as Volvox spheres composed of thousands of biflagellate cells, illustrate early steps toward multicellularity through coordinated cell division and differentiation.^[8] Filamentous types, exemplified by Spirogyra, form unbranched chains of cylindrical cells in freshwater environments, while multicellular seaweeds like Ulva (sea lettuce) develop macroscopic, sheet-like thalli up to several meters in size in marine settings. Green algae represent a paraphyletic group, meaning they do not form a single evolutionary clade but are instead split between two major lineages within Viridiplantae: the Chlorophyta and the Streptophyta (the latter including charophyte green algae and land plants).^[8] This paraphyly arises because land plants evolved from within the streptophyte algae, leaving the remaining green algae as a grade rather than a clade.^[8] Key subgroups highlight this diversity; prasinophytes, early-diverging unicellular members of Chlorophyta, exhibit primitive flagellated forms with scaly surfaces.^[9] The core chlorophytes comprise a wide array of mostly aquatic species, including both marine and freshwater inhabitants with varied reproductive strategies.^[6] In contrast, charophytes within Streptophyta, such as stoneworts (Chara) and conjugating algae (Spirogyra), are predominantly freshwater forms and phylogenetically closest to land plants, sharing traits like phragmoplast-mediated cell division.^[9]

Importance and Distribution

Green algae serve as primary producers in aquatic ecosystems, forming the base of food webs and supporting diverse marine and freshwater communities through photosynthesis. They contribute significantly to global oxygen production as part of phytoplankton assemblages, with marine species playing a key role in atmospheric oxygenation.^[10]^[11] Additionally, certain green algae, such as those in the genus Ulva, act as indicators of water quality; their prolific blooms often signal nutrient pollution from eutrophication, leading to green tides that disrupt coastal habitats.^[12]^[13] In scientific research, green algae are valued as model organisms for investigating fundamental biological processes. The unicellular alga Chlamydomonas reinhardtii, for instance, is widely used to study photosynthesis, chloroplast function, cell motility via flagella, and genetic mechanisms due to its simple structure and ease of genetic manipulation.^[14]^[15] Volvocine green algae, including Chlamydomonas and multicellular forms like Volvox, further aid in exploring evolutionary transitions in multicellularity and biofluid dynamics.^[16] Green algae exhibit a broad global distribution, inhabiting freshwater, marine, and terrestrial environments, with approximately 90% of species favoring freshwater habitats. They thrive in diverse settings, from coastal seas to soil surfaces, and extend to extreme conditions such as polar snowfields, where species like Chlamydomonas nivalis cause characteristic red snow and contribute to primary production in cold ecosystems.^[17]^[18]^[19] Symbiotically, they form mutualistic associations with fungi in lichens, where genera like Trebouxia provide photosynthetic products to the fungal partner, enabling lichen survival in harsh terrestrial niches; about 50-70% of lichens rely on such green algal symbionts from the Trebouxiaceae family.^[20] In marine contexts, green algae such as zoochlorellae serve as symbiotic partners in some sea anemones and other cnidarians on reefs, supplying nutrients via photosynthesis.^[21]

Cellular and Structural Biology

Cell Structure and Ultrastructure

Green algal cells exhibit a eukaryotic organization typical of photosynthetic protists, featuring a plasma membrane, nucleus, mitochondria, and chloroplasts derived from a primary endosymbiosis event. The overall cell morphology varies widely, from unicellular flagellates to colonial or multicellular forms, but ultrastructural features provide key diagnostic traits for the group.^[22] The cell wall of green algae is primarily composed of cellulose microfibrils embedded in a matrix of polysaccharides and proteins, providing structural support and protection. In chlorophytes, such as those in the Chlorophyceae, cell walls often include hydroxyproline-rich glycoproteins, which contribute to flexibility and adhesion in species like Chlamydomonas. In contrast, charophytes incorporate more complex components, including sporopollenin—a durable, hydrophobic polymer akin to that in land plant spores—enhancing resistance to environmental stresses in taxa like Coleochaete.^[1]^[23] Chloroplasts in green algae are bounded by two membranes and contain thylakoids that are frequently stacked into grana, facilitating efficient light harvesting similar to those in land plants. A prominent feature is the pyrenoid, a dense, proteinaceous structure within the chloroplast stroma that serves as a site for starch accumulation and carbon concentrating mechanisms. In motile green algae, an eyespot or stigma—a lipid droplet layer associated with the chloroplast—enables phototaxis by directing light to underlying photoreceptors.^[24]^[25]^[26] Motile cells in many green algae, exemplified by Chlamydomonas reinhardtii, possess a flagellar apparatus consisting of two anteriorly inserted flagella emerging from basal bodies. These flagella exhibit a typical 9+2 axoneme structure and are adorned with mastigonemes—fine, hair-like appendages composed of hydroxyproline-rich glycoproteins that influence swimming behavior and hydrodynamic properties.^[27] Mitotic division in green algae displays significant variation reflective of phylogenetic divisions. In the Chlorophyceae, mitosis is typically closed, with the nuclear envelope remaining intact throughout division, and a persistent intranuclear spindle facilitating chromosome segregation. Conversely, charophytes undergo open mitosis, involving nuclear envelope breakdown and formation of a phragmoplast—a microtubule array that guides cell plate deposition—mirroring the process in land plants.^[28]^[22] Unlike many other algal groups that store photosynthetic products as floridean starch or laminarin in the cytoplasm, green algae accumulate starch granules directly within the chloroplast, often surrounding the pyrenoid, which distinguishes them biochemically and links them evolutionarily to embryophytes.^[29]

Pigments and Photosynthetic Apparatus

Green algae primarily contain chlorophyll a and chlorophyll b as their main photosynthetic pigments, which enable the absorption of light in the blue and red regions of the visible spectrum. Chlorophyll a exhibits absorption maxima at approximately 430 nm and 662 nm, while chlorophyll b absorbs strongly at 453 nm and 642 nm, broadening the range of wavelengths utilized for photosynthesis compared to organisms lacking chlorophyll b. These pigments are embedded within the thylakoid membranes of chloroplasts, where they drive the light-dependent reactions.^[30]^[31] Accessory pigments such as β-carotene and various xanthophylls, including lutein, complement the chlorophylls by absorbing light in the green-yellow spectrum and transferring energy to the reaction centers. The photosynthetic apparatus in green algae features photosystems I and II (PSI and PSII), which are multi-subunit complexes located in the thylakoid membranes and function similarly to those in land plants. Associated with these photosystems are light-harvesting complexes (LHCs), particularly the LHCII trimer for PSII and LHCI for PSI, which enhance energy capture and transfer efficiency through antenna pigments arranged in a way that minimizes energy loss.^[32]^[33]^[34] The plastid genome of green algae encodes essential core proteins for the photosynthetic apparatus, such as subunits of PSI, PSII, and the cytochrome b₆f complex, though many genes have undergone transfer to the nuclear genome in more derived lineages, necessitating import of nuclear-encoded proteins back into the plastid. This nuclear-to-plastid gene transfer reflects evolutionary adaptations for coordinated regulation of photosynthesis.^[35]^[36] Variations in pigment composition occur across green algal groups; for instance, some prasinophytes possess prasinoxanthin, a distinctive xanthophyll that serves as a chemotaxonomic marker and contributes to their light-harvesting capabilities.^[37] Overall, the pigment profile—dominated by chlorophylls reflecting green light—imparts the characteristic coloration to green algae. The quantum yield of oxygen evolution in green algae, measuring photosynthetic efficiency, is comparable to that of higher plants, typically ranging from 0.10 to 0.12 molecules of O₂ per quantum of light absorbed under optimal conditions.^[32]

Reproduction and Life Cycles

Asexual Reproduction

Asexual reproduction in green algae enables rapid clonal propagation without genetic recombination, facilitating population expansion in favorable environments and survival under stress. This process encompasses various mechanisms adapted to the diverse morphologies of green algae, from unicellular to multicellular forms, and predominates in many species, particularly those with haplontic life cycles where the dominant phase is haploid.^[38] In unicellular green algae such as Chlamydomonas, asexual reproduction occurs through multiple fission, a modified mitotic process where a mother cell undergoes successive nuclear divisions followed by cytokinesis, producing 2 to 16 daughter cells known as autospores. These autospores are released upon rupture of the mother cell wall and develop into new vegetative cells, allowing for efficient proliferation under nutrient-rich conditions. This mechanism contrasts with simple binary fission seen in some prokaryotes but is optimized for the eukaryotic cell cycle in Chlamydomonas reinhardtii.^[39]^[38] Filamentous green algae, exemplified by Spirogyra, reproduce asexually via fragmentation, where the elongated filament breaks into segments due to mechanical stress or environmental factors, and each fragment regenerates into a complete new filament through apical cell growth. This vegetative propagation is straightforward and supports colonization of freshwater habitats, with fragments capable of rapid re-establishment without specialized structures. Akin to fragmentation, some species form akinetes or other resting fragments for dormancy.^[40]^[41] Zoospore formation represents a dispersive asexual strategy in more complex green algae like Ulva, where specialized sporangia develop within the thallus and release biflagellate zoospores equipped with flagella for motility and dispersal in aquatic environments. These motile spores settle and germinate into new thalli, promoting spread in marine settings and responding to environmental cues such as light and nutrients for synchronous release. In contrast, aplanospores serve as non-motile asexual propagules produced in response to adverse conditions, such as desiccation or nutrient limitation; the protoplast contracts within a thickened cell wall, forming a resistant spore that germinates when conditions improve, as observed in charophyte green algae.^[42]^[43] Parthenogenesis occurs in certain green algae, including some ulvophytes, where unfertilized eggs or gametes develop directly into new individuals, maintaining haploid lineages in haplontic life cycles without meiosis or fusion. This form of apomixis enhances reproductive assurance in sparse populations, as seen in Ulva prolifera where parthenogenetic zoospores contribute to bloom formation.^[44]

Sexual Reproduction and Pheromones

Sexual reproduction in green algae encompasses a range of strategies that promote genetic recombination through gamete fusion, varying from primitive isogamy to advanced oogamy. Recent studies (as of 2023) have identified key genes that regulate the development of female and male reproductive cells in volvocine algae, providing insights into the evolution of sexual differentiation.^[45] Isogamy involves the fusion of morphologically similar, flagellated gametes of equal size, as observed in species like Chlamydomonas reinhardtii, where plus and minus mating types pair based on molecular recognition.^[46] Anisogamy features gametes of differing sizes but similar motility, an intermediate form seen in some volvocine algae, bridging isogamy and more differentiated systems.^[47] Oogamy represents the most derived type, with small, flagellated sperm fertilizing large, non-motile eggs; this occurs in filamentous genera such as Oedogonium, where dwarf male filaments produce sperm that swim to oogonia containing eggs.^[48] Life cycles in green algae typically follow haplontic or haplobiontic patterns, differing in the dominance of haploid versus alternating generations. In the haplontic cycle, prevalent in conjugating forms like Spirogyra, the multicellular organism is haploid, and meiosis occurs zygotically in the diploid zygote, which germinates directly into new haploid filaments.^[49] Haplobiontic cycles feature isomorphic alternation of generations, as in Ulva species, where morphologically identical haploid gametophytes produce flagellated gametes that fuse to form a diploid sporophyte, which then undergoes meiosis to release haploid zoospores.^[50] A distinctive sexual process in zygnematophytes, such as Spirogyra, is conjugation, where non-flagellated, amoeboid gametes fuse without motility. Adjacent cells from two filaments form a thin cytoplasmic bridge or conjugation tube, through which protoplasts migrate and merge, often in a scalariform (ladder-like) or lateral manner, leading to zygote formation within one cell.^[51] This flagella-independent mechanism suits freshwater environments with low flow, enabling efficient gamete contact.^[52] Sex pheromones play a crucial role in coordinating mating, particularly in volvocine green algae like Volvox carteri, where a glycoprotein pheromone at concentrations below 10^{-16} M induces sexual differentiation, gamete release, and aggregation in a species-specific manner. These chemical signals ensure compatible mating types interact, enhancing reproductive success by triggering developmental shifts from asexual to sexual phases.^[53] Following fusion, the zygote develops into a thick-walled zygospore, serving as a dormant stage resistant to environmental stresses. In species like Spirogyra and Chlamydomonas, zygospores feature multi-layered walls with sporopollenin, enabling survival during unfavorable conditions such as desiccation or cold, before meiosis and germination resume the life cycle.^[54] This dormancy facilitates overwintering and dispersal, contributing to the ecological persistence of green algae.^[55]

Taxonomy and Phylogeny

Historical Classification

The classification of green algae began with Carl Linnaeus in his 1753 work Species Plantarum, where algae were placed within the artificial class Cryptogamia, encompassing plants without conspicuous seeds or flowers. Green algae were primarily aggregated under the genus Conferva, a broad category that included diverse filamentous and branched forms such as Cladophora and Spirogyra, reflecting the era's reliance on gross morphology rather than pigmentation or cellular details. This grouping treated Conferva as a catch-all for non-vascular, aquatic plants with thread-like structures, though Linnaeus also recognized separate genera like Ulva for sheet-like forms and Chara for more complex, stonewort-like algae.^[56]^[57] In the 19th century, Swedish phycologist Carl Adolph Agardh advanced algal taxonomy through a more systematic approach based on pigmentation, thallus form, and reproductive characteristics. In his 1817 Synopsis Algarum Scandinaviae, Agardh divided algae into sections by color, placing green algae under Ulvoideae, which encompassed about 45 genera distinguished by their chlorophyll content and simple thalli. By the 1820s and 1830s, Agardh formalized the class Chlorophyceae (initially termed Chlorospermeae) for green algae, emphasizing their uniform green pigmentation due to chlorophyll a and b, alongside starch storage and varied thallus types from unicellular to multicellular. This system marked a shift toward natural classification, though it still lumped disparate forms together based on superficial traits.^[58] Twentieth-century classifications increasingly emphasized reproductive structures, cytology, and life cycle details to refine groupings. Systems like that of H. Kylin in the 1920s and 1930s separated green algae into distinct classes—Chlorophyceae for flagellate, unicellular, and colonial forms with isogamous or anisogamous reproduction; Ulvophyceae for siphonous, coenocytic thalli with advanced vegetative propagation; and Charophyceae for branched, calcified forms with oogamous sexual reproduction—highlighting differences in gamete motility and zygote retention. F. E. Fritsch's influential 1935 treatise The Structure and Reproduction of the Algae standardized these classes within Chlorophyta, prioritizing microscopic features like chloroplast arrangement, pyrenoid presence, and flagellar orientation to delineate subgroups, while treating Chlorophyceae as the core class encompassing most freshwater and terrestrial green algae. This cytology-based framework became a benchmark for pre-molecular taxonomy.^[59] By the 1960s, accumulating evidence from electron microscopy and comparative morphology revealed the paraphyletic nature of traditional Chlorophyta groupings, as certain green algal lineages (notably Charophyceae) shared advanced reproductive and structural traits with embryophytes (land plants), sparking debates on whether to subsumed land plants within an expanded Chlorophyta or restrict the division to non-terrestrial forms. These challenges underscored the limitations of purely morphological systems, setting the stage for later revisions without resolving the inclusion of embryophytes.^[60]^[61]

Modern Phylogenetic Classification

The modern phylogenetic classification of green algae, part of the Viridiplantae clade, recognizes two primary lineages: the Chlorophyta, encompassing the core green algae, and the Streptophyta, which includes charophyte algae and land plants. This division is supported by extensive molecular data, resolving the paraphyletic nature of traditional green algae groupings. Analyses of nuclear, plastid, and mitochondrial genomes have established these lineages as monophyletic, with Chlorophyta diversifying primarily in marine and freshwater environments, while Streptophyta are predominantly freshwater inhabitants.^[62] A basal clade, Prasinodermophyta, was formally recognized in 2020 as a distinct phylum sister to the Chlorophyta-Streptophyta split, comprising unicellular and macroscopic forms adapted to low-light, oligotrophic conditions. This clade includes classes such as Prasinodermophyceae (e.g., Prasinococcus, a picoplankter) and Palmophyllophyceae (e.g., Palmophyllum, deep-water macroalgae), featuring unique carbon-concentrating mechanisms and dual NAD+ biosynthesis pathways. Prasinodermophyta diverged early, with genomic similarities of approximately 38.5% to Chlorophyta and 33.9% to Streptophyta, highlighting its foundational role in green plant evolution.^[63] Within Chlorophyta, multi-gene phylogenies using up to 2,698 nuclear genes across 70 species have delineated major clades, including the UTC assemblage (Ulvophyceae, Trebouxiophyceae, Chlorophyceae) and basal prasinophytes. The UTC clade is characterized by phycoplast-mediated cytokinesis and diverse morphologies, with Ulvophyceae exhibiting non-monophyly (split into Ulvophyceae I and II, the latter sister to Chlorophyceae), Trebouxiophyceae often terrestrial or symbiotic, and Chlorophyceae featuring motile flagellated cells. Prasinophytes form a polyphyletic grade of early-diverging lineages, totaling 7-9 classes in Chlorophyta schemes (e.g., Mamiellophyceae, Nephroselmidophyceae, Chloropicophyceae), with scaly flagella in many and high oceanic biodiversity in orders like Mamiellales, which dominate picophytoplankton communities.^[64]^[62] Genomic studies from the 2020s, incorporating 557 single-copy orthologous genes and plastid datasets (74 genes across 63 taxa), have further clarified prasinophyte polyphyly by reassigning groups like Palmophyllophyceae to Prasinodermophyta and confirming deep relationships via maximum-likelihood and Bayesian inference. These approaches, building on 18S rRNA and plastid gene analyses, have resolved longstanding ambiguities in clade boundaries, emphasizing convergent evolution in traits like flagellar scales across prasinophyte lineages.^[63]^[62]

Relationship to Land Plants

Green algae, particularly the charophyte lineages within the Streptophyta clade, share a close evolutionary relationship with land plants (Embryophyta), forming a monophyletic group that diverged from other green algae (Chlorophyta) over 500 million years ago.^[65] This clade unites diverse charophyte groups, such as the Charales (stoneworts) and Zygnematales (conjugating algae), with embryophytes, supported by molecular phylogenetic analyses that resolve Streptophyta as the sister lineage to Chlorophyta.^[66] The shared ancestry is evidenced by ultrastructural and biochemical similarities that predate the transition to terrestrial habitats, indicating that key adaptations for plant-like development originated in aquatic algal ancestors.^[67] Several shared innovations highlight this phylogenetic proximity, including phragmoplast-mediated cytokinesis, where a microtubule-based phragmoplast guides the formation of a cell plate during cell division, a process characteristic of advanced charophytes and all land plants but absent in chlorophytes.^[68] Additionally, both groups possess rosette-shaped cellulose-synthesizing complexes composed of cellulose synthase A (CesA) proteins embedded in the plasma membrane, which assemble linear cellulose microfibrils essential for cell wall integrity; this complex evolved near the base of charophyte algae and is retained in embryophytes.^[69] In advanced charophytes like those in the Charales and Coleochaetales, sperm ultrastructure shows striking similarities to that in basal land plants, featuring flagellated, biflagellate sperm with comparable acronematic morphology and mitochondrial arrangement, facilitating motile fertilization in aquatic environments.^[70] Phylogenetic studies from 2020, incorporating nuclear and organellar genomes, have refined the closest algal relatives of land plants to the Zygnematophyceae, a class of conjugating green algae that lack flagella and exhibit non-motile gametes, challenging earlier models favoring flagellate groups like Charales.^[71] This sister-group relationship is robust across multiple analyses, despite the apparent simplicity of zygnematophyte morphology, and underscores genomic adaptations for terrestrial transition that likely arose after divergence.^[72] Furthermore, evidence of horizontal gene transfer (HGT) from bacterial sources into streptophyte algal ancestors has contributed to land plant evolution, with transferred genes such as those encoding aquaporins and other transporters exapted for drought tolerance functions in embryophytes.^[73] The fossil record provides temporal context for this relationship, with charophyte-like fossils appearing in the Ordovician period (~453–449 million years ago), predating the earliest unequivocal land plant fossils from the Silurian (~430 million years ago) and supporting an aquatic origin for the streptophyte lineage before embryophyte colonization of land.^[74] These Ordovician marine charophyte gyrogonites and vegetative remains, such as Tarimochara miraclensis, indicate that charophytes had diversified in shallow marine environments well before the adaptive radiation of terrestrial plants.^[75]

Evolution

Origins and Endosymbiosis

The photosynthetic capabilities of green algae originated through a primary endosymbiosis event approximately 1.5 billion years ago, in which a heterotrophic eukaryotic host engulfed and retained a free-living cyanobacterium as an endosymbiont. This ancient merger established the plastids characteristic of the Viridiplantae lineage, which encompasses green algae and land plants, enabling oxygenic photosynthesis within eukaryotic cells. The event is considered part of a singular primary endosymbiosis that also gave rise to the Archaeplastida supergroup, uniting the green (Viridiplantae), red (Rhodophyta), and glaucophyte lineages through a shared cyanobacterial ancestor.^[76]^[77]^[78] Following endosymbiosis, extensive gene transfer from the cyanobacterial endosymbiont to the host nucleus—known as endosymbiotic gene transfer (EGT)—facilitated the integration of the organelle and its metabolic pathways into the eukaryotic cell. This process relocated hundreds of genes to the nucleus, allowing nuclear control over plastid functions while leaving only essential genes in the organelle. As a result, plastid genomes in green algae have been dramatically reduced in size, typically ranging from 100 to 200 kilobase pairs and encoding 100-200 genes, a fraction of the original cyanobacterial genome.^[79]^[76] The timing of this foundational event is supported by both fossil and molecular evidence, though the fossil record provides a minimum age constraint. The earliest unambiguous fossils of photosynthetic eukaryotes with primary plastids are represented by Bangiomorpha pubescens, a red alga from the Archaeplastida, dated to approximately 1.05 billion years ago, indicating that crown-group diversification had already begun by the Mesoproterozoic. However, Bayesian molecular clock analyses, calibrated with fossil constraints, place the origin of Archaeplastida and primary endosymbiosis between 1.5 and 1.8 billion years ago, during the Paleoproterozoic, highlighting a significant temporal gap between molecular estimates and the preserved fossil record.^[80] Unlike certain algal groups such as euglenophytes or some dinoflagellates, green algae retain primary plastids derived directly from the cyanobacterial endosymbiont and have not undergone secondary endosymbiosis, preserving a simpler double-membrane structure around their chloroplasts.^[77]

Evolutionary Diversification

The green algal lineage originated from a small, unicellular, marine biflagellate ancestor approximately 1 billion years ago during the Mesoproterozoic era. This ancestral form, resembling modern prasinophytes in its simple morphology, underwent early diversification within the prasinophyte grade, establishing the basal branches of the Chlorophyta clade and setting the stage for broader adaptive radiations. Fossil evidence from this period, including Proterozoic microfossils such as Proterocladus antiquus, corroborates the presence of these early unicellular forms, with branched filaments up to 2 mm long preserved in Chinese Tonian deposits.^[22] Following this basal phase, the Chlorophyta radiated prominently into marine planktonic niches, where lineages like the Mamiellophyceae achieved ecological dominance, comprising up to 55% of green microalgal operational taxonomic units in global ocean sampling efforts and contributing significantly to picophytoplankton biomass. In parallel, the Streptophyta diverged toward freshwater environments, with charophyte algae evolving greater structural complexity, including macroscopic thalli in orders like the Charales that feature whorled branches and calcified reproductive structures. These habitat shifts highlight a core dichotomy in green algal evolution: oceanic adaptation in chlorophytes versus continental freshwater colonization in streptophytes.^[81]^[82]^[83] Key innovations further drove diversification within these clades, such as the secondary loss of flagella in the Zygnematophyceae, which eliminated flagellated stages and centrioles, enabling efficient sexual reproduction through conjugation in non-motile, filamentous or coccoid forms. Similarly, the Volvocales exhibited the emergence of coloniality, with species forming organized cell aggregates that represent a transitional step toward differentiated multicellularity, as seen in genera like Gonium and Pandorina. Paleozoic fossils, including Devonian charophyte fructifications from around 400 million years ago and widespread gyrogonites in Carboniferous limestones, document this increasing complexity in streptophyte lineages.^[84]^[85]^[86]^[87] Environmental drivers, including Neoproterozoic oxygenation events and enhanced nutrient availability in expanding shallow marine and freshwater systems, profoundly shaped these radiations by facilitating planktonic expansions and habitat transitions from marine to limnic settings. These factors promoted the proliferation of photosynthetic niches, underscoring the interplay between geochemical changes and algal adaptive evolution.^[88]

Physiology and Biochemistry

Metabolic Processes

Green algae primarily employ the Calvin-Benson cycle for carbon dioxide fixation, where the enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) catalyzes the reaction of CO₂ with ribulose-1,5-bisphosphate to produce two molecules of 3-phosphoglycerate, establishing the predominant C3 photosynthetic pathway unlike the C4 mechanism observed in certain land plants.^[89] This cycle operates within the plastid stroma, integrating with light-dependent reactions to convert inorganic carbon into organic compounds essential for growth and biomass accumulation.^[90] Following carbon fixation, green algae synthesize starch as their primary carbohydrate storage reserve in the plastids through an ADP-glucose-dependent pathway, involving enzymes such as ADP-glucose pyrophosphorylase and starch synthase, which polymerize glucose units into amylose and amylopectin granules.^[91] This process mirrors the prokaryotic origins of their plastids and serves as a key energy depot, mobilized during periods of darkness or stress to support metabolic demands.^[92] For energy metabolism, green algae conduct glycolysis in the cytosol to break down glucose into pyruvate, generating ATP and NADH, while the tricarboxylic acid (TCA) cycle in the mitochondria oxidizes acetyl-CoA to produce additional reducing equivalents for oxidative phosphorylation under aerobic conditions.^[93] Under anaerobic conditions, such as in oxygen-deprived environments, species like Chlamydomonas reinhardtii shift to fermentation, producing ethanol via pyruvate decarboxylase and alcohol dehydrogenase to regenerate NAD⁺ and maintain glycolytic flux.^[94] Nitrogen assimilation in green algae begins with nitrate uptake and reduction by nitrate reductase to nitrite in the cytosol, followed by nitrite reductase converting it to ammonium, which is then incorporated into glutamine via glutamine synthetase and glutamate synthase in the GS/GOGAT pathway.^[95] Some chlorophytes exhibit remnants of urea cycle enzymes, such as arginase, enabling partial cycling for nitrogen recycling under specific conditions, though a complete cycle is absent compared to animals or certain other algae.^[96] Lipid metabolism in green algae facilitates energy storage through the accumulation of triacylglycerols (TAGs) in lipid bodies, particularly under nutrient stress like nitrogen limitation, where carbon flux is redirected from growth to neutral lipid synthesis via Kennedy pathway enzymes in the endoplasmic reticulum, highlighting their potential as biofuel feedstocks.^[97] This stress-induced TAG buildup can exceed 50% of dry biomass in species like Chlorella vulgaris, balancing energy reserves with environmental adaptability.^[98]

Environmental Physiology

Green algae exhibit remarkable physiological adaptations to abiotic stresses, enabling survival in diverse environments ranging from hypersaline waters to extreme temperatures. These adaptations involve intricate molecular and cellular mechanisms that maintain cellular homeostasis under fluctuating conditions. Key responses include osmoregulation to counter salinity changes, photoprotection against excess light, thermal acclimation to temperature extremes, and detoxification of heavy metals through chelating compounds. Osmoregulation in green algae is primarily achieved through the accumulation of organic osmolytes and regulation of ion channels to counteract osmotic stress in hypersaline conditions. For instance, the halotolerant microalga Dunaliella salina synthesizes and accumulates glycerol as a compatible solute, which balances external osmotic pressure without disrupting cellular metabolism, allowing growth in salinities up to 5 M NaCl.^[99] This process involves rapid activation of glycerol-3-phosphate dehydrogenase upon salinity shock, leading to intracellular glycerol concentrations exceeding 50% of cell dry weight. Ion channels, such as voltage-gated K⁺ and Cl⁻ channels, further facilitate turgor regulation by enabling selective ion fluxes across the plasma membrane.^[99] Under high light irradiance, green algae employ photoprotective mechanisms to dissipate excess energy and prevent photooxidative damage to photosystem II. Non-photochemical quenching (NPQ) is a primary strategy, where absorbed light energy is converted to heat rather than being used in photochemistry, particularly through pH-dependent activation in the thylakoid lumen. In species like Chlamydomonas reinhardtii, NPQ can quench up to 80% of excess excitation energy during sudden light increases.^[100] Complementing NPQ, carotenoid shifts occur, with an increase in protective carotenoids such as zeaxanthin and β-carotene under high irradiance; for example, exposure to intensities above 1000 µmol photons m⁻² s⁻¹ induces a 2- to 3-fold rise in β-carotene content in Haematococcus pluvialis, enhancing antioxidant capacity.^[101] Temperature responses in green algae involve the induction of stress proteins to stabilize cellular structures during thermal extremes. In Chlamydomonas reinhardtii, heat shock above 35°C triggers the synthesis of heat shock proteins (HSPs), such as HSP70 and small HSPs, which act as molecular chaperones to refold denatured proteins and prevent aggregation, allowing survival up to 42°C for short periods.^[102] For cold acclimation, snow algae like Chlamydomonas nivalis produce antifreeze-like proteins, including polyproline type II helical structures, which inhibit ice crystal growth by binding to ice surfaces and lowering the freezing point by 1-2°C, facilitating persistence in subzero environments.^[103] These proteins are upregulated during cold exposure below 4°C, contributing to membrane fluidity maintenance and photosynthetic efficiency.^[104] Heavy metal detoxification in green algae relies on the biosynthesis of phytochelatins, cysteine-rich peptides that chelate toxic metals like cadmium and mercury, sequestering them in vacuoles to mitigate oxidative stress. In Chlorella vulgaris, exposure to 10 µM Cd²⁺ induces phytochelatin synthase activity, producing PC₂ and PC₃ isoforms that bind up to 90% of intracellular metal ions within hours.^[105] This mechanism enhances bioremediation potential, as metal-laden algal biomass can accumulate concentrations 10-100 times higher than surrounding media, supporting applications in wastewater treatment.^[105]

Ecology and Applications

Habitats and Ecological Roles

Green algae occupy diverse habitats, ranging from marine and freshwater environments to terrestrial settings. In oceanic ecosystems, planktonic forms such as prasinophytes serve as key primary producers, contributing significantly to marine primary production through their role in the photosynthetic picoeukaryote community.^[106] Benthic species like Cladophora thrive in freshwater systems, attaching to hard substrates in the littoral zones of rivers and lakes where they form dense mats in nutrient-enriched waters. Terrestrial green algae, including aero-terrestrial species, colonize moist soils, rocks, and tree bark, aiding in soil stabilization and weathering processes in arid or semi-arid regions.^[107] Symbiotic associations enhance the ecological integration of green algae. Approximately 90% of lichen species host green algae as photobionts, providing photosynthetic support to the fungal partner in nutrient-poor environments.^[108] Additionally, certain green algae form intracellular mutualisms with freshwater invertebrates, such as sponges in the genus Ephydatia, where the algae supply photosynthates in exchange for protection and nutrients.^[109] In trophic dynamics, green algae form the foundational layer of aquatic food webs, serving as primary producers that sustain herbivores like zooplankton and benthic grazers, thereby supporting higher trophic levels including fish populations.^[110] However, excessive growth can lead to harmful algal blooms; for instance, Ulva species proliferate in eutrophic coastal waters, outcompeting other organisms and causing oxygen depletion upon decay.^[111] Green algae contribute to biogeochemical cycles through carbon sequestration, as sunken biomass from blooms, such as those of Ulva, accumulates in marine sediments, locking away organic carbon over long timescales.^[112] Some free-living forms participate in nitrogen dynamics via associations with nitrogen-fixing bacteria, enhancing nutrient availability in oligotrophic habitats, though direct fixation is rare among green algae themselves.^[113] Responses to global environmental changes are pronounced, with studies from the 2020s indicating that warming and ocean acidification promote increased bloom frequency and intensity. For example, elevated temperatures and CO₂ levels have been shown to accelerate growth and nutrient uptake in bloom-forming species like Ulva lactuca, exacerbating eutrophication risks in coastal systems.^[114]^[115]

Human Uses and Economic Importance

Green algae have significant applications in biotechnology, particularly as sources of nutraceuticals and aquaculture feeds. Species such as Chlorella vulgaris are cultivated for their rich content of proteins, vitamins (including B12 and D), polyunsaturated fatty acids, and antioxidants, which are incorporated into dietary supplements promoting human health benefits like immune support and detoxification.^[116]^[117] Similarly, Haematococcus pluvialis serves as the primary commercial source of natural astaxanthin, a potent antioxidant comprising 3-7% of its dry weight, used in nutraceuticals for anti-inflammatory and skin health applications; industrial production employs two-stage cultivation in photobioreactors to optimize yields.^[118]^[119] In aquaculture, green microalgae like Chlorella provide high-nutritional-value feeds for larval fish, shrimp, and bivalves, offering essential amino acids, lipids, and pigments that enhance growth and survival rates while reducing reliance on fishmeal.^[120]^[121] In biofuel production, lipid-rich chlorophytes such as Chlamydomonas reinhardtii are engineered for biodiesel via genetic modifications that boost triacylglycerol accumulation. Overexpression of key enzymes like chloroplast glyceraldehyde-3-phosphate dehydrogenase has increased lipid yields by redirecting carbon flux, with 2020s advancements in CRISPR-based editing enabling targeted enhancements for higher productivity and scalability.^[122]^[123] These efforts address economic viability by improving biomass output under stress conditions like nitrogen limitation. Green algae contribute to bioremediation by absorbing heavy metals and treating wastewater. Scenedesmus species, for instance, bioaccumulate up to 87% of cadmium, 85% of chromium, and 90% of iron from contaminated effluents through biosorption and intracellular uptake mechanisms.^[124] Additionally, Scenedesmus obliquus and related strains remove hexavalent chromium via metabolic processes, while their nutrient uptake capabilities—enhanced by CO2 supplementation—facilitate phosphorus and nitrogen removal in wastewater, supporting integrated lipid production for dual environmental and economic benefits.^[125]^[126] As research models, green algae enable studies on fundamental biological processes. Volvox carteri exemplifies multicellularity evolution, with its developmental genetics—featuring somatic-germ cell differentiation—providing insights into embryogenesis and morphogenesis in volvocine algae.^[128]^[129] Genomic sequencing of Prasinodermophyta, such as the 2020 assembly of Prasinoderma coloniale (a ~25 Mb genome encoding 7,139 genes), reveals early green lineage innovations, furnishing resources for synthetic biology applications like pathway engineering in related chlorophytes.^[63] Green algae, such as Chlorella vulgaris, function as biofertilizers to enhance soil health. Extracts supply organic matter, phytohormones, and nutrients that improve soil structure, water retention, and microbial activity, promoting crop growth as sustainable alternatives to synthetic fertilizers.^[130] As of 2025, research on algal microbiomes highlights their role in enhancing resilience to environmental stressors, advancing sustainable biotechnology applications.^[131] Emerging uses include large-scale cultures for climate mitigation, where microalgae capture CO2 at rates up to 1.5 kg/m²/year through photosynthesis, converting it to biomass for sequestration or biofuels, thereby offsetting emissions in integrated systems.^[132]^[133]

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Plastidic starch synthesis in green algae and plants occurs via ADP-glucose in likeness to pro- karyotes from which plastids have evolved.
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Metabolic Constraints for Carbon Partitioning between Oil and Starch
Under normal photoheterotrophic conditions, acetate (C2) is assimilated to form acetyl-CoA and feeds into the TCA cycle and respiration or into the glyoxylate ...
[93]
Fermentation metabolism and its evolution in algae - PMC
For Chlamydomonas, dark fermentation leads to the production of formate, acetate, and ethanol in a 2:1:1 ratio (Mus et al., 2007; Figure 2).
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Understanding nitrate assimilation and its regulation in microalgae
Nitrate assimilation is a key process for nitrogen (N) acquisition in green microalgae. Among Chlorophyte algae, Chlamydomonas reinhardtii has resulted to ...
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Evolution and metabolic significance of the urea cycle in ... - PubMed
May 12, 2011 · Here we show that the exosymbiont-derived ornithine-urea cycle, which is similar to that of metazoans but is absent in green algae and plants, facilitates ...Missing: remnants chlorophytes
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Metabolic regulation of triacylglycerol accumulation in the green algae
Jan 23, 2016 · The great need for more sustainable alternatives to fossil fuels has increased our research interests in algal biofuels.
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Enhancing microalgal lipid accumulation for biofuel production
The major challenge of enhancing microalgal lipid accumulation lies in overcoming the trade-off between microalgal cell growth and lipid accumulation.
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Characterization of the mechanism of prolonged adaptation ... - Nature
Sep 19, 2016 · For these microorganisms, salinity and osmotic stress tolerance are prerequisites for survival. To date, two general adaptive strategies to ...
[99]
Non-photochemical Quenching: From Light Perception to ...
Jan 8, 2022 · Plants and microalgae have developed photoprotective mechanisms collectively referred to as non-photochemical quenching to minimize such potential damage.
[100]
Microalgae for the production of lipid and carotenoids: a review with ...
Oct 4, 2018 · Microalgae can overproduce lipids or carotenoids under stress condition such as high salt, high light, or nutrient limitation [11, 12]. For ...
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The Chlamydomonas heat stress response - Wiley Online Library
Mar 6, 2015 · We found HSP genes to be induced when Chlamydomonas cells were shifted from 25°C to at least 37°C and cells to tolerate a maximum temperature ...Summary · Why Study the Heat Stress... · Heat Stress Signal Transduction
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Polyproline type II helical antifreeze proteins are widespread in ...
Jun 1, 2023 · Antifreeze proteins (AFPs) bind to ice crystals to prevent organisms from freezing. A diversity of AFP folds has been found in fish and ...
[103]
Cold Adaptation Mechanisms of a Snow Alga Chlamydomonas ...
Chlamydomonas nivalis UTEX 2824 is a snow alga isolated from the Sierra Nevada Mountains in California that has been used as a research model to study the ...
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(PDF) Action Potential in Charophytes - ResearchGate
Aug 6, 2025 · Studies of action potentials in the vacuole and cytoplasm of Nitella. ... cytoplasmic streaming induced by membrane excitation in characean ...
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A physical perspective on cytoplasmic streaming - PMC
Two spiralling bands of molecular motors at the cell periphery drive the cellular fluid up and down at speeds up to 100 µm s−1, motion that has been ...
[106]
Microalgal Metallothioneins and Phytochelatins and Their Potential ...
Eukaryotic microalgae appear to be particularly suitable to use in bioremediation since they require only sunlight and inorganic nutrients for their growth, can ...
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Evidence-based green algal genomics reveals marine diversity and ...
Mar 31, 2016 · Prasinophytes are a major lineage of unicellular green algae that can contribute significantly to marine primary production [2–4]. In the ...
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A critical review of soil algae as a crucial soil biological component ...
Mar 19, 2024 · Aero-terrestrial algae are ecologically and economically valuable bioresources contributing to carbon sequestration, sustenance of soil health, and fertility.Soil Algae Of Extreme... · Algae Of Forest Soils · Soil Algal Diversity And Its...
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The Lichens' Microbiota, Still a Mystery? - PMC - PubMed Central
Mar 30, 2021 · The large majority of the approximately 20.000 described lichen species are formed with a green alga as photobiont, whereas only 10% of lichens ...
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Freshwater sponge hosts and their green algae symbionts - NIH
In many freshwater habitats, green algae form intracellular symbioses with a variety of heterotrophic host taxa including several species of freshwater sponge.
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Aquatic food webs | National Oceanic and Atmospheric Administration
Sep 30, 2025 · Producers like plants and algae are at the base of the food web. ... All herbivores are primary consumers that eat from the base of the food chain ...
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Macroalgal blooms caused by marine nutrient changes resulting ...
Mar 4, 2020 · Ulva is the most common macrofouling algae because it can adhere to marine equipment such as hulls and docks that are periodically submerged by ...Macroalgal Blooms Caused By... · 2 Ulva And Green Tides In... · 5 Ulva Sporulation...
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Fate and carbon sequestration potential of sunken macroalgae in ...
The massive sunken macroalgae following green tides holds substantial potential for ocean carbon sequestration, providing new insights into the carbon sequ.Results · Microbial Metabolic... · Discussion And Conclusions<|separator|>
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Ecosystem consequences of a nitrogen-fixing proto-organelle - PMC
Recent studies demonstrate endosymbiosis of N-fixing bacteria into algal hosts, but the ecological significance of such partnerships remains unexplored. In a ...
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Species-Specific Responses of Bloom-Forming Algae to the Ocean ...
The results indicate that the concurrent rises in temperature and pCO 2 level significantly boosted the growth and nutrient uptake rates of U. lactuca.Missing: 2020s | Show results with:2020s
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Climate Change and Harmful Algal Blooms: Insights and perspective
Oct 28, 2025 · Climate change is transforming aquatic ecosystems. Coastal waters have experienced progressive warming, acidification, and deoxygenation that will intensify ...Missing: 2020s | Show results with:2020s
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Potential of Chlorella as a Dietary Supplement to Promote Human ...
Chlorella products contain numerous nutrients and vitamins, including D and B 12 , that are absent in plant-derived food sources.
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Chemical Compounds, Bioactivities, and Applications of Chlorella ...
This review presents the chemical composition, bioactive properties, and diverse applications of Chlorella vulgaris, a green microalga widely recognized for ...2.5. Vitamins And Minerals · 3.1. Antioxidant Activity · 3.9. Detoxification And...
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Astaxanthin-Producing Green Microalga Haematococcus pluvialis
Among the commercially important microalgae, Haematococcus pluvialis is the richest source of natural astaxanthin which is considered as “super anti-oxidant.” ...
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From present to prosperity: assessing the current status and ...
Dec 15, 2023 · Haematococcus pluvialis is a promising source of natural astaxanthin (3.00–7.00% by dry weight) which is being globally cultivated at an ...
[121]
Potential of microalgae as a sustainable feed ingredient for ...
Nov 20, 2021 · This review paper will explore the potential of producing microalgae biomass as an ingredient of aquaculture feed.
[122]
Use of Algae in Aquaculture: A Review - MDPI
Micro- and macroalgae used as a feed supplement have demonstrated positive effects on the green water culture production of various planktivorous species, ...
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Chlamydomonas as a model for biofuels and bio-products production
Chlamydomonas reinhardtii's ease of transformation, large toolset, and natural variety has led to strains with significantly increased biofuels and bio-product ...Missing: 2020s | Show results with:2020s
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Increasing lipid production in Chlamydomonas reinhardtii through ...
Apr 18, 2024 · This study aimed to boost lipid production in C. reinhardtii by overexpressing chloroplast glyceraldehyde-3-phosphate dehydrogenase (GAPDH), a key enzyme in ...Missing: 2020s | Show results with:2020s
[125]
Scenedesmus sp. Harvesting by Using Natural Coagulant after ...
This study indicated that Scenedesmus sp. can remove heavy metals cadmium (87.24%), chromium (85.55%), and ferum (90.35%), respectively.Scenedesmus Sp. Harvesting... · 3. Results And Discussion · 3.1. Heavy Metals Removal By...
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Utilization of the microalga Scenedesmus quadricauda for ...
The purpose of this study was to evaluate the bioremediation potential of the microalga Scenedesmus quadricauda in removing hexavalent chromium (Cr (VI)) ...2. Materials And Methods · 3. 3. Results And Discussion · 3.1. Biomass Growth Under...
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Microalgae in removing heavy metals from wastewater
This review highlights current advances in using different species of microalgae to treat various industrial effluents and mechanism of removal of heavy metals ...
[128]
Influence of organic matter and CO 2 supply on bioremediation of ...
Oct 30, 2019 · This research evaluated the influence of organic matter (OM) and CO 2 addition on the bioremediation potential of two microalgae typically used for wastewater ...
[129]
Origins of multicellular complexity: Volvox and the volvocine algae
Volvox and related algae have become an important model system for the major transition from unicellular to multicellular life.
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Volvox: A simple algal model for embryogenesis, morphogenesis ...
Nov 1, 2016 · The multicellular green alga Volvox carteri (Volvox) is an excellent model for the investigation of developmental mechanisms and their evolutionary origins.
[131]
Algae as Bio-fertilizers: Between current situation and future ...
Mar 19, 2022 · This review provides an effective approach to increase soil fertility via environmentally friendly bio-based fertilizer using micro and macro algae.Missing: charophyte | Show results with:charophyte
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The potential microalgae-based strategy for attaining carbon ...
Previous research indicates that microalgae can fix CO2 at rates exceeding 1.5 kg/m2/year under optimal conditions, and produce lipids with high content of ...
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Minimizing carbon footprint via microalgae as a biological capture
However, microalgae have emerged as a bio-sequestration of carbon dioxide, in which carbon dioxide is buried into biomass via photosynthesis, and the harvested ...