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Symbiogenesis

Symbiogenesis is the evolutionary process by which new species, tissues, organs, or even entire lineages emerge through the stable integration and coevolution of symbiotic organisms, often via endosymbiosis, where one organism lives inside another.^[1] This mechanism contrasts with traditional views of evolution driven solely by gradual mutations, emphasizing cooperation and genomic merging as drivers of major innovations, such as the origin of eukaryotic cells from prokaryotic symbionts.^[2] The concept traces its roots to early 20th-century botanists like Konstantin Mereschkowski, who in 1905 proposed symbiotic origins for chloroplasts, but it gained prominence through the work of Lynn Margulis, who in 1967 articulated the serial endosymbiosis theory in her paper "On the Origin of Mitosing Cells."^[3] Margulis expanded this in her 1970 book Origin of Eukaryotic Cells, arguing that mitochondria originated from engulfed α-proteobacteria capable of aerobic respiration, chloroplasts from cyanobacteria enabling photosynthesis, and even eukaryotic flagella from spirochete bacteria, though the latter lacks genomic support.^[2] These ideas faced initial skepticism but were bolstered by accumulating evidence, including phylogenetic analyses showing mitochondrial and plastid genomes closely resembling those of their bacterial counterparts.^[2] Symbiogenesis extends beyond organelles to broader evolutionary patterns, such as the diversification of multicellular life forms and the Proterozoic fossil record, where symbiotic integrations likely fueled the rise of complex protists and animals.^[1] Modern consensus holds that eukaryotic evolution involved a foundational symbiosis between an archaeal host and a bacterial endosymbiont (the mitochondrion), with subsequent acquisitions like plastids in plants, underscoring symbiosis as a recurrent force in generating biodiversity. Recent examples include the 2024 identification of the nitroplast in marine algae, a nitrogen-fixing endosymbiont evolving into an organelle.^[4]^[5] This process highlights how horizontal gene transfer and metabolic cooperation, rather than competition alone, have shaped life's major transitions.^[4]

Historical Development

Early Proposals

The concept of symbiogenesis emerged in the late 19th and early 20th centuries amid a scientific landscape influenced by pre-Darwinian vitalism, which emphasized life's inherent complexity and organizational principles beyond mere mechanical processes, and early microscopy that revealed enigmatic intracellular structures in eukaryotic cells. Botanists and microscopists observed phenomena such as the independent division of organelles like chloroplasts, which appeared autonomous yet integrated within host cells, prompting speculative ideas about symbiotic origins rather than de novo creation. These observations, facilitated by improved light microscopes, highlighted similarities between intracellular bodies and free-living microbes, setting the stage for theories challenging traditional views of cellular evolution.^[6] In 1883, German botanist Andreas Schimper proposed one of the earliest ideas of symbiotic origins for chloroplasts while studying plant cell development. Observing the division and inheritance of chlorophyll granules under the microscope, Schimper suggested that these structures—now known as plastids—arose from the symbiotic association of distinct organisms, specifically likening them to engulfed cyanobacteria capable of independent reproduction. His hypothesis posited that photosynthetic eukaryotes resulted from the fusion of a heterotrophic host with a photosynthetic symbiont, though he provided no experimental mechanisms for integration.^[6]^[7] Building on Schimper's work, Russian botanist Konstantin Mereschkowski advanced the symbiogenesis theory in 1905, explicitly proposing that chloroplasts originated from symbiotic cyanobacteria. In his paper "Über Natur und Ursprung der Chromatophoren in den Pflanzen," Mereschkowski argued that the chromatophores (chloroplasts) of plants were not evolved de novo but represented captured, blue-green algae (cyanobacteria) that retained reproductive autonomy within a heterotrophic host cell. He extended this in 1910 to suggest a "theory of two plasmas," where the eukaryotic cell arose from multiple symbiotic unions, including one forming the nucleus, drawing on vitalist notions of life's modular assembly.^[8]^[6] French physiologist Paul Portier contributed to the discourse in his 1918 book Les Symbiotes, proposing intracellular symbiosis as a universal principle of life and specifically identifying mitochondria as derived from symbiotic bacteria. Portier envisioned all cells as composite entities formed by interlocked symbionts, with mitochondria functioning as aerobic bacteria that enhanced host metabolism through endosymbiotic integration, observed via early staining techniques that revealed their rod-like forms. His ideas, rooted in microbiological observations, extended symbiogenesis beyond plants to animal and fungal cells.^[9]^[6] These early proposals, while visionary, faced significant limitations due to the era's technological constraints, including the absence of genetic evidence to confirm organelle autonomy or bacterial ancestry, and the inability to culture isolated organelles, which led many to dismiss them as speculative artifacts of microscopy rather than viable evolutionary mechanisms. Lacking molecular tools, proponents relied on morphological analogies, which were insufficient against prevailing reductionist paradigms favoring autogenous origins for organelles. These ideas remained marginal until later synthesized by Lynn Margulis in the 1960s.^[8]^[7]

Modern Advancements

In the early 20th century, American biologist Ivan Wallin advanced symbiogenesis by conducting experiments in the 1920s and 1930s to culture mitochondria as independent bacterial entities outside host cells, though these efforts ultimately failed due to contamination issues and methodological limitations.^[10] His 1927 book Symbionticism and the Origin of Species synthesized these attempts, arguing that mitochondria originated from symbiotic bacteria and extending the idea to broader evolutionary implications for speciation, influencing later theorists despite contemporary dismissal.^[11] The theory gained renewed prominence in 1967 through Lynn Margulis's (then Lynn Sagan) seminal paper "On the Origin of Mitosing Cells," which formalized the serial endosymbiosis theory (SET) by proposing that mitochondria derived from engulfed alphaproteobacteria and chloroplasts from cyanobacteria, marking a key refinement of earlier symbiogenetic ideas.^[12] Margulis expanded this framework in her 1970 book Origin of Eukaryotic Cells, integrating ultrastructural, biochemical, and phylogenetic evidence to argue for a symbiotic origin of eukaryotic organelles, emphasizing serial acquisitions that built cellular complexity.^[13] These works shifted symbiogenesis from fringe speculation to a testable hypothesis, building briefly on pre-20th-century proposals like those of Konstantin Mereschkowski while incorporating modern microscopy and microbiology. Acceptance of SET accelerated post-1970 due to molecular discoveries in the 1960s, particularly the identification of DNA within mitochondria by Margit M.K. Nass and Sylvan Nass in 1963, who observed DNase-sensitive fibers via electron microscopy, confirming organelles' semi-autonomous genetic systems akin to bacteria. Similar findings for chloroplast DNA further supported endosymbiotic origins, providing empirical validation that propelled symbiogenesis into mainstream evolutionary biology by the 1980s.^[14] Thomas Cavalier-Smith contributed significantly to 20th- and 21st-century refinements by integrating membrane biogenesis with symbiogenesis, proposing in his 2000 paper that membrane heredity—via symbiont-derived lipid synthesis—drove early chloroplast evolution and eukaryotic compartmentalization.^[15] His models also emphasized archaeal host integration, suggesting that the eukaryotic nuclear membrane arose from archaeal plasma infoldings during symbiosis, reconciling symbiogenesis with archaeal-bacterial chimerism in a 2010 synthesis. Recent phylogenomic studies, including analyses as of 2023, have pinpointed mitochondrial ancestors more precisely within the Pelagibacterales order of alphaproteobacteria, with genomic analyses of the endosymbiont Odyssella thessalonicensis reinforcing shared ancestry with free-living Pelagibacter ubique and the mitochondrion of the eukaryote Reclinomonas americana^[16], highlighting streamlined marine bacteria as the likely progenitors. These phylogenomic studies, leveraging expanded bacterial sampling, refine SET by underscoring the role of ecologically dominant, low-energy-adapted symbionts in eukaryotic emergence.^[17]

Core Mechanisms

Engulfment and Symbiotic Integration

Symbiogenesis begins with the phagocytosis-like engulfment of free-living prokaryotes by a host cell, typically envisioned as an archaeon from the Asgard superphylum.^[18] In this process, the host cell, likely an H₂-dependent heterotroph, captures an alphaproteobacterium (for mitochondria) or cyanobacterium (for plastids) through membrane invagination, similar to modern phagocytic mechanisms observed in some bacteria and early eukaryotes. This engulfment forms a food vacuole around the endosymbiont, preventing immediate digestion and allowing initial survival within the host cytosol. Evidence from phylogenomic analyses supports the archaeal host's role, highlighting shared informational genes with eukaryotes while metabolic genes derive from the bacterial endosymbiont.^[19] Following engulfment, the relationship evolves through the establishment of mutual benefits, transitioning the endosymbiont from a potential prey or parasite to a symbiotic partner. The endosymbiont provides the host with enhanced energy production, such as ATP export via adenine nucleotide translocators in the case of mitochondrial precursors, in exchange for protection from external threats and access to host-derived metabolites like carbon sources and inorganic ions. This metabolic complementarity is evident in models where the alphaproteobacterial endosymbiont oxidizes host-produced H₂ or fermentative byproducts, yielding energy that boosts host fitness in oxygenated environments. Over generations, these exchanges stabilize the association, with the host gaining respiratory efficiency and the endosymbiont benefiting from a nutrient-rich, sheltered niche.^[20]^[19] Membrane dynamics during integration result in the characteristic double-membrane structure of organelles. The inner membrane originates from the endosymbiont's plasma membrane, retaining bacterial lipid composition and respiratory complexes, while the outer membrane derives from the host's phagocytic vacuole membrane, which fuses or modifies post-engulfment to encase the endosymbiont. This topology is supported by the presence of bacterial-derived porins like VDAC in the outer membrane and the impermeability of the inner membrane to protons, essential for energy generation. These membranes facilitate controlled exchange, with the outer membrane allowing diffusion of small molecules and the inner maintaining proton gradients for ATP synthesis.^[21] The evolution of protein import systems, such as the TOM (translocase of the outer membrane) and TIM (translocase of the inner membrane) complexes in mitochondria, enables endosymbiont-derived proteins encoded by the host nucleus to be targeted back into the organelle. TOM complexes, featuring β-barrel channels like Tom40 of bacterial origin, recognize mitochondrial targeting signals on precursor proteins in the cytosol and translocate them across the outer membrane. TIM complexes, including Tim23 for presequence pathways, then guide proteins through the inner membrane, often powered by electrochemical gradients. These systems arose convergently post-engulfment, adapting bacterial export machinery and host components to support organelle autonomy while integrating with host cellular processes.^[22] The stages of symbiotic integration progress from initial parasitic or commensal interactions to stable mutualism across multiple generations. Early phases involve sporadic engulfment events where the endosymbiont may exploit host resources without reciprocity, but selective pressures favor hosts retaining viable symbionts that provide metabolic advantages. Gradual physiological coupling, such as synchronized division and membrane contact sites, reinforces dependency, culminating in obligate mutualism where neither can survive independently. This progression is inferred from comparative genomics and experimental models of bacterial endosymbioses, emphasizing metabolic interdependence over time. Gene transfer later stabilizes this integration, but the core physical and physiological symbiosis establishes the foundation.^[19]

Endosymbiotic Gene Transfer

Endosymbiotic gene transfer (EGT) refers to the relocation of genetic material from the genomes of endosymbiotic organelles, such as mitochondria and plastids, to the host cell's nucleus, a process central to the integration of these organelles into eukaryotic cells. This transfer reduces the autonomy of the endosymbiont-derived organelles while enabling the host nucleus to control their functions through nuclear-encoded proteins that are imported back into the organelles. EGT is an ongoing phenomenon, with evidence of recent transfers detectable in modern genomes, and it has profoundly shaped eukaryotic evolution by creating chimeric nuclear genomes enriched with prokaryotic genes.^[23] The primary mechanisms of EGT involve the release and integration of organelle DNA into the nuclear genome. Direct transfer often occurs through the lysis of organelles, which liberates DNA fragments that can be captured and incorporated into nuclear chromosomes via non-homologous end-joining or microhomology-mediated repair. For instance, in plants with multiple chloroplasts per cell, lysis facilitates the release of plastid DNA, enabling its uptake and integration, as demonstrated experimentally in tobacco where large chunks of chloroplast DNA (up to 131 kb) were transferred to the nucleus. Vesicle-mediated transfer has also been proposed, where DNA is packaged in vesicles budding from organelle membranes and transported to the nucleus, though this mechanism is less frequently documented and may complement lysis in certain contexts. Additionally, transposon activity can promote integration by mobilizing DNA fragments within the nucleus, facilitating the stable insertion of endosymbiont-derived sequences, as seen in cases where retrotransposition aids the domestication of transferred genes.^[24]^[24]^[25] Through EGT, more than 90% of ancestral mitochondrial and plastid genes have been lost or relocated to the nucleus across eukaryotic lineages, resulting in highly reduced organelle genomes.^[26] This massive gene relocation is evident in the human mitochondrial genome, which retains only 13 protein-coding genes out of an estimated 1,000 in the original alphaproteobacterial endosymbiont. For plastids, transfers have integrated thousands of cyanobacterial genes into the nucleus, with about 18% of nuclear genes in model plants like Arabidopsis thaliana tracing back to the plastid ancestor. A notable example is the bacterial RNA polymerase subunit gene rpoB, originally mitochondrial or plastidial, which in many eukaryotes is now nuclear-encoded and expressed as a precursor protein targeted back to the organelle. These transfers underscore the selective pressures favoring nuclear control, where genes for metabolic and housekeeping functions are preferentially relocated while a core set remains in organelles for rapid local regulation.^[23]^[27]^[23] To function in their original organelles after transfer, relocated genes acquire nuclear targeting signals, primarily N-terminal transit peptides that direct proteins through organelle import machinery. These amphipathic peptides, typically 20-80 amino acids long, evolve rapidly post-transfer and interact with specific translocases like TOC (translocon at the outer chloroplast membrane) or TOM (mitochondrial outer membrane) complexes for import. The evolution of such targeting sequences is crucial for EGT success, as without them, transferred gene products would accumulate in the cytosol rather than restoring organelle function. Experimental reconstructions in yeast and plants confirm that transit peptide addition enables functional complementation of organelle deficiencies.^[28]^[29] The consequences of EGT include dramatic organelle genome reduction, leaving mitochondria and plastids with compact genomes encoding only essential, hydrophobic, or redox-sensitive proteins. In humans, the mitochondrial genome encodes just 13 proteins involved in the electron transport chain, necessitating nuclear oversight for the remaining ~1,400 mitochondrial proteins. Similarly, plastid genomes retain ~100-200 genes, primarily for photosynthesis and translation, while the nucleus manages the bulk of organelle proteome assembly. This dependency enhances host control but introduces vulnerabilities, such as reliance on import systems, and contributes to chimeric nuclear genomes that complicate phylogenetic reconstructions. Overall, EGT exemplifies how endosymbiosis drives genomic innovation, with initial engulfment providing the prerequisite for sustained gene flow.^[23]^[24]^[30]

Primary Endosymbiosis

Protomitochondrial Origin

The protomitochondrial origin refers to the endosymbiotic event in which an alphaproteobacterial symbiont was engulfed by an archaeal host cell, leading to the establishment of mitochondria as essential organelles in eukaryotic cells. Phylogenetic analyses indicate that the host was likely an archaeon closely related to the Asgard superphylum, which possesses genomic features suggestive of early eukaryotic traits such as actin-like proteins and membrane remodeling capabilities.^[31] This host may have exhibited hydrogen-dependent metabolism, facilitating a syntrophic relationship with the symbiont.^[32] The symbiont is identified as an alphaproteobacterium, with molecular evidence pointing to a free-living lineage, particularly relatives among marine bacteria in the Iodidimonadales order.^[33] This event is estimated to have occurred approximately 1.5 to 2 billion years ago, following the Great Oxidation Event (GOE) around 2.4 billion years ago, which increased atmospheric oxygen levels and enabled the evolution of aerobic processes.^[34] The timing aligns with fossil and molecular clock data supporting the emergence of eukaryotic complexity in the Paleoproterozoic era, though estimates vary between 1.45 and 1.8 billion years ago.^[35] A pivotal innovation of this symbiosis was the adoption of oxidative phosphorylation by the protomitochondrion, allowing efficient aerobic respiration and ATP production under oxygenated conditions.^[20] The protomitochondria supplied ATP to the host via mechanisms such as ADP/ATP carriers, reducing the host's reliance on fermentation and enabling energy-intensive processes that promoted larger cell sizes and increased metabolic complexity.^[20] This integration marked a foundational step in eukaryogenesis, with subsequent endosymbiotic gene transfer further stabilizing the organelle's role.^[33]

Cyanobacterial Plastid Origin

The primary endosymbiosis leading to the origin of plastids involved the engulfment of a photosynthetic cyanobacterium by a heterotrophic eukaryotic host cell that had already acquired mitochondria through an earlier endosymbiotic event.^[36] This host was likely an archaeal-derived eukaryote capable of phagocytosis, providing the cellular machinery for symbiotic integration.^[37] The cyanobacterial symbiont belonged to a lineage closely related to modern unicellular cyanobacteria, such as those in the Synechococcus-Prochlorococcus clade or early-branching forms like Gloeomargarita lithophora, which possessed the genetic toolkit for oxygenic photosynthesis including photosystems I and II.^[38]^[39] This endosymbiotic event is dated to approximately 1 to 1.5 billion years ago, based on molecular clock analyses of nuclear and organellar genes, as well as fossil evidence of early photosynthetic eukaryotes, though estimates range from 0.85 to 1.6 billion years ago.^[36]^[40]^[41] The timeline places it well after the mitochondrial endosymbiosis, around 1.5 to 2 billion years ago, allowing the host to benefit from the symbiont's photosynthetic capabilities for energy production beyond respiration. Over time, massive endosymbiotic gene transfer from the cyanobacterial genome to the host nucleus reduced the organelle's genome to a compact plastome, while retaining key photosynthetic genes.^[42] A defining feature of these primary plastids is their retention of two bounding membranes: an inner membrane derived from the cyanobacterium's plasma membrane and an outer membrane from its cell wall, which facilitate protein import via TOC/TIC translocons.^[43] Thylakoid membranes, critical for light-dependent reactions of photosynthesis, originated from the symbiont's internal invaginations and lipid phase transitions involving galactolipids like monogalactosyldiacylglycerol, evolving independently of the host's endomembrane system.^[41]^[44] This structural simplicity contrasts with more complex organelles but underscores the direct inheritance of cyanobacterial photosynthetic machinery. The resulting plastids are distributed exclusively within the Plantae supergroup, encompassing glaucophytes (e.g., Cyanophora), red algae (Rhodophyta), green algae (Chlorophyta), and land plants (Embryophyta), all sharing a monophyletic origin from this single endosymbiotic event.^[45] Phylogenetic analyses of plastid-targeted proteins and ribosomal RNA confirm this unity, with no evidence of additional primary plastid acquisitions in other eukaryotic lineages.^[46] This restricted distribution highlights the event's pivotal role in establishing photosynthetic autotrophy in eukaryotes.

Organelle Derivatives and Evolution

Endomembrane System and Nucleus

The nuclear envelope, a defining feature of eukaryotic cells, is thought to have arisen through invaginations of the host cell's plasma membrane, forming a double-membrane barrier that encloses the genome and facilitates selective transport via nuclear pore complexes.^[47] This structure likely emerged as an adaptation to compartmentalize genetic material in the context of increasing cellular complexity during early eukaryogenesis.^[48] Protomitochondrial interactions may have played a role in stabilizing this envelope, with mitochondrial-derived vesicles potentially contributing to membrane remodeling and integration shortly after endosymbiosis.^[49] The endomembrane system, encompassing the endoplasmic reticulum (ER) and Golgi apparatus, primarily derives from the archaeal host's membrane architecture but was significantly enhanced by endosymbiont-derived components following mitochondrial acquisition.^[37] Bacterial lipids from the protomitochondrion, such as ester-linked phospholipids, replaced archaeal ether lipids, enabling greater membrane fluidity and the formation of dynamic vesicular structures essential for protein and lipid processing.^[49] Additionally, proteins encoded by genes transferred from the endosymbiont to the host nucleus supported the expansion of trafficking machinery, including those involved in lipid synthesis and membrane curvature.^[49] The symbiotic integration of the protomitochondrion imposed selective pressures that drove the evolution of sophisticated membrane trafficking, as the host cell required mechanisms to engulf, retain, and exchange materials with the endosymbiont while managing metabolic byproducts.^[49] This necessity likely accelerated the development of the endomembrane network to handle increased vesicular transport demands, transforming a simple prokaryotic-like membrane system into the compartmentalized eukaryotic one.^[50] Key evidence for these endosymbiotic influences includes the conservation of SNARE proteins across the endomembrane system and in organelle interactions, which mediate specific vesicle fusion events and are absent in prokaryotes, underscoring their emergence as a eukaryotic innovation post-endosymbiosis.^[51] Endosymbiotic gene transfer further provided nuclear-encoded membrane proteins that integrated into host-derived systems, reinforcing the interconnected evolution of these compartments.^[37]

Hydrogenosomes and Mitosomes

Hydrogenosomes and mitosomes represent highly reduced, anaerobic derivatives of mitochondria that have adapted to oxygen-poor environments through extensive gene loss and functional specialization. These organelles, collectively known as mitochondrion-related organelles (MROs), illustrate the evolutionary plasticity of the mitochondrial lineage following its initial endosymbiotic integration. While retaining core import machinery for proteins, they have diverged from canonical mitochondria by eliminating oxidative phosphorylation and repurposing for alternative metabolic roles.^[20] Hydrogenosomes are double-membrane-bound organelles found in anaerobic protists such as trichomonads, including the human pathogen Trichomonas vaginalis. Unlike mitochondria, they lack cristae and a genome, instead generating ATP via substrate-level phosphorylation in a fermentative pathway. Pyruvate is metabolized by pyruvate:ferredoxin oxidoreductase (PFOR) to acetyl-CoA, with electrons transferred to ferredoxin and ultimately to hydrogenase, producing molecular hydrogen (H₂) as a byproduct. This process supports energy production in oxygen-deprived niches, such as the vertebrate urogenital tract, and highlights hydrogenosomes' role in anaerobic carbohydrate catabolism.^[52]^[53] Mitosomes, identified in diverse anaerobic and microaerophilic eukaryotes including microsporidians like Encephalitozoon cuniculi and Trachipleistophora hominis, are even more streamlined organelles that have completely lost energy-producing capabilities. They lack both a genome and respiratory functions, serving primarily as sites for iron-sulfur (Fe-S) cluster biogenesis essential for cytosolic and nuclear proteins. Key components, such as the scaffold protein Isu1, cysteine desulfurase Nfs1, and frataxin, localize to mitosomes and facilitate Fe-S cluster assembly, a remnant of the ancient mitochondrial machinery conserved across eukaryotes. This function underscores mitosomes' indispensability despite their minimal proteome, preventing reliance on cytosolic alternatives for Fe-S maturation.^[54]^[55] Both hydrogenosomes and mitosomes trace their ancestry to an alphaproteobacterial endosymbiont that gave rise to all mitochondria, with divergence occurring after the initial symbiogenetic event through lineage-specific gene loss and endosymbiotic gene transfer to the host nucleus. Phylogenetic analyses of conserved proteins, such as heat shock proteins Hsp60 and Hsp70, place hydrogenosomal and mitosomal homologs within the mitochondrial clade, confirming a shared origin rather than independent acquisitions. This evolutionary trajectory involved the complete elimination of organellar DNA in mitosomes and the absence or extreme reduction of genomes in hydrogenosomes, rendering them fully dependent on nuclear-encoded, imported proteins for maintenance.^[56]^[20]

Nitroplasts

Nitroplasts represent a recently discovered endosymbiotic organelle specialized for nitrogen fixation, marking the first known example of such an organelle in eukaryotes.^[5] This breakthrough came in 2024 through studies on the marine haptophyte alga Braarudosphaera bigelowii, which harbors the cyanobacterial symbiont UCYN-A2 as an integrated cellular component.^[5] Researchers confirmed its organelle status by demonstrating tight integration into the host cell's architecture, synchronized division with the host, and import of host-encoded proteins essential for its function, distinguishing it from a mere endosymbiont.^[5] The nitroplast originates from UCYN-A, a unicellular nitrogen-fixing cyanobacterium that has undergone extensive integration with its eukaryotic host.^[5] Unlike typical cyanobacteria, UCYN-A lacks the capacity for oxygenic photosynthesis due to the absence of photosystem II and associated genes.^[5] Instead, it expresses nitrogenase enzymes to fix atmospheric nitrogen (N₂) into bioavailable forms, providing a critical nutrient to the host alga in nutrient-poor marine environments.^[5] The symbiont's genome is highly reduced, measuring approximately 1.5 megabases (Mb), reflecting metabolic streamlining and dependence on the host for amino acids, nucleotides, vitamins, and other essentials.^[57] Structurally, the nitroplast is enveloped by the host's endoplasmic reticulum (ER), facilitating protein import and nutrient exchange while protecting the oxygen-sensitive nitrogenase from cellular respiration.^[5] This symbiosis holds profound evolutionary implications, potentially representing a third primary endosymbiosis event in eukaryotes alongside mitochondria and plastids.^[5] Molecular clock analyses date the establishment of the UCYN-A association to the late Cretaceous period, approximately 100 million years ago, making the nitroplast a relatively young organelle compared to ancient ones like chloroplasts.^[5] The ongoing endosymbiotic gene transfer and protein targeting observed suggest it is at an early stage of organelle evolution, offering a unique window into the mechanisms driving symbiont-to-organelle transitions.^[5] Nitroplasts are distributed specifically within certain haptophyte lineages, primarily Braarudosphaera bigelowii and related species, where the UCYN-A2 sublineage predominates; other UCYN-A sublineages associate with different haptophytes as endosymbionts without confirmed full organelle status as of 2024.^[5]^[58] This association is globally prevalent in oligotrophic oceans, contributing significantly to marine nitrogen budgets, but remains confined to these algal groups without evidence of broader horizontal spread.^[5]

Organellar Genomes

Mitogenomes and Plastomes

Mitochondrial genomes, or mitogenomes, are typically compact, circular DNA molecules inherited maternally in most eukaryotes. In animals, mitogenomes range from 16 to 18 kilobases (kb) in length and encode 13 core protein-coding genes essential for the oxidative phosphorylation system, along with 22 transfer RNAs (tRNAs) and 2 ribosomal RNAs (rRNAs), totaling 37 genes.^[59] In plants, mitogenomes are significantly larger, often exceeding 300 kb and reaching up to approximately 1 megabase (Mb) due to the incorporation of non-coding sequences and additional genes, encoding around 30-40 proteins, numerous tRNAs, and rRNAs.^[60] These genomes reflect the bacterial ancestry of mitochondria, with much of the original genetic content having been transferred to the nucleus over evolutionary time.^[61] Plastid genomes, known as plastomes, are also predominantly circular in primary endosymbionts such as those in land plants and green algae, though linear forms occur in certain lineages like some red algae. Typical plastomes measure 120-200 kb and contain approximately 100-200 genes, including those for ribosomal components (rRNAs and tRNAs), RNA polymerase subunits, and a suite of proteins dedicated to photosynthesis, such as the psbA gene encoding the D1 protein of photosystem II.^[62]^[63] In land plants, plastomes often feature a quadripartite structure with two inverted repeats flanking large and small single-copy regions, enhancing stability and gene dosage.^[64] Both mitogenomes and plastomes retain bacterial-like features, including the organization of genes into operons and polycistronic transcription, where multiple genes are transcribed as a single precursor mRNA that is subsequently processed.^[65] This prokaryotic-style expression facilitates coordinated production of organelle components. Variations exist, such as in ciliates, where mitogenomes are linear and fragmented into multiple chromosomes, with genes often split and requiring trans-splicing for maturation.^[66]

Reduced and Non-Photosynthetic Genomes

Non-photosynthetic plastids, derived from ancient cyanobacterial endosymbionts, exhibit significant genome reduction compared to their photosynthetic counterparts, which typically range from 120 to 160 kb in size. In organisms such as apicomplexan parasites, the apicoplast—a non-photosynthetic plastid—harbors a highly compact circular genome of approximately 35 kb that encodes around 30 genes primarily involved in housekeeping functions like transcription, translation, and essential metabolic processes, including support for peptidoglycan synthesis required for organelle division.^[67]^[68] These genes enable the apicoplast to perform non-photosynthetic roles, such as isoprenoid biosynthesis, which are critical for parasite survival.^[69] Examples of such reduced genomes abound in diverse eukaryotic lineages, including non-photosynthetic algae and parasitic plants. In heterotrophic stramenopiles such as Spumella (chrysophyte) and Pteridomonas (dictyochophycean) species, leucoplast-like plastids retain compact genomes of 33–63 kb, having lost all photosynthetic genes while preserving those for tRNA-mediated heme biosynthesis and iron-sulfur cluster assembly.^[70]^[71] Similarly, in mycoheterotrophic plants such as Monotropa uniflora, the plastid genome is reduced to approximately 27 kb, retaining a minimal set of genes for ribosomal proteins and RNA components to support metabolic functions like fatty acid synthesis, despite the complete absence of photosynthesis-related machinery.^[72]^[73] These organelles, often referred to as leucoplasts in plant contexts or chromeroplasts in certain algal relatives, underscore the plasticity of plastid evolution, where gene loss is offset by nuclear-encoded imports.^[74] Extreme cases of genome minimization highlight the limits of organelle dependency. For instance, the plastid genome of the holoparasitic plant Pilostyles aethiopica measures just 11.3 kb, encoding only five or six potentially functional genes focused on core translational components, representing one of the most reduced plastomes known among land plants.^[75] More recently, as of 2024, the plastid genome of the non-photosynthetic stramenopile Leucomyxa sphaerocephala was sequenced at approximately 20 kb, retaining a reduced gene set for essential functions like translation while exemplifying further compaction.^[76] This drastic shrinkage reflects the evolutionary pressures favoring efficiency in non-autonomous organelles, where most former plastid functions have been transferred to the host nucleus. The primary driver of such reductions is endosymbiotic gene transfer (EGT), whereby genes from the endosymbiont's genome are relocated to the host nucleus, enforcing biochemical dependency and streamlining the organelle for specialized, non-redundant roles.^[77] This process, coupled with relaxed selection on photosynthesis, promotes the loss of unnecessary genes while retaining a core set essential for organelle maintenance, ultimately enhancing host control over the endosymbiont.^[78]

Supporting Evidence

Molecular and Structural Evidence

Mitochondria and chloroplasts replicate through binary fission, a process analogous to bacterial cell division, where the organelles constrict and divide using ring-like structures involving proteins such as FtsZ in chloroplasts, which is a bacterial homolog.^[79] This division mechanism ensures equitable distribution during host cell mitosis and underscores the prokaryotic heritage of these organelles. In contrast, eukaryotic cytosolic division relies on microtubules, highlighting the distinct, bacteria-derived autonomy of organelle proliferation.^[80] Organelles possess 70S ribosomes, smaller and structurally similar to those in bacteria, enabling independent protein synthesis sensitive to antibiotics that target prokaryotic ribosomes, such as chloramphenicol.^[19] These ribosomes translate organelle-encoded genes, including components of the electron transport chain in mitochondria and photosynthetic apparatus in chloroplasts, further evidencing their bacterial ancestry.^[81] The ribosomal RNA sequences in these organelles cluster phylogenetically with bacterial counterparts, reinforcing endosymbiotic origins.^[19] Shared lipid compositions provide additional molecular evidence; for instance, cardiolipin, a diphosphatidylglycerol lipid unique to bacterial plasma membranes, predominates in the inner mitochondrial membrane, stabilizing respiratory complexes and absent from other eukaryotic membranes.^[82] Cardiolipin biosynthesis in mitochondria involves bacterial-type synthases, linking organelle membrane biogenesis to prokaryotic pathways.^[83] Similarly, the outer membranes of mitochondria and chloroplasts contain porins—β-barrel proteins forming aqueous channels for metabolite diffusion—that are homologous to bacterial outer membrane porins, facilitating passive transport akin to Gram-negative bacteria.^[84] The double-membrane architecture of mitochondria and chloroplasts represents a structural hallmark of endosymbiosis, with the inner membrane derived from the bacterial plasma membrane and the outer from the host's phagosomal vesicle.^[37] Protein import into these organelles occurs via translocon complexes; in mitochondria, the TOM (translocase of the outer membrane) and TIM (translocase of the inner membrane) systems incorporate components with bacterial homologs, such as chaperone-like proteins aiding precursor translocation reminiscent of bacterial secretion machinery.^[85] Phylogenetic analyses show that many organelle-resident proteins, including those in metabolic pathways, cluster with alphaproteobacterial (for mitochondria) or cyanobacterial (for chloroplasts) sequences, supporting their endosymbiotic integration.^[19]

Phylogenetic and Fossil Evidence

Phylogenetic analyses of mitochondrial genes consistently place their origin within the Alphaproteobacteria, a diverse bacterial group, supporting the endosymbiotic acquisition of mitochondria from a free-living alphaproteobacterial ancestor.^[86] Similarly, genes encoded in plastids and nuclear genes of plastid origin cluster robustly with those from cyanobacteria, particularly non-marine lineages, confirming the cyanobacterial provenance of primary plastids in the Archaeplastida supergroup.^[87] The eukaryotic nuclear genome exhibits a chimeric composition, with genes involved in information processing—such as replication, transcription, and translation—predominantly tracing to archaeal ancestors, while those related to energy metabolism and operational functions are largely bacterial in origin, reflecting the archaeal-bacterial fusion central to eukaryogenesis.^[88] This dichotomy underscores the host's archaeal heritage and the endosymbiont's contribution to metabolic innovations like oxidative phosphorylation. Molecular clock analyses, calibrated with fossil constraints, estimate the mitochondrial endosymbiosis at approximately 2 billion years ago (Ga), aligning with the emergence of the eukaryotic domain during the Paleoproterozoic.^[89] Molecular clock analyses estimate primary plastid endosymbiosis between approximately 0.9 and 2.1 billion years ago (Ga), with many studies suggesting around 1.2 Ga.^[35]^[90] Fossil records provide corroborative timelines for symbiogenesis. Microfossils from the ~1.88 Ga Gunflint Chert in Ontario, Canada, include acritarch-like structures that some studies propose as possible early eukaryotic fossils with intracellular features, though their interpretation as organelle precursors remains controversial, suggesting potential mitochondrion-bearing eukaryotes by the late Paleoproterozoic.^[91] Evidence for photosynthetic eukaryotes appears later, with Bangiomorpha pubescens from ~1.05 Ga deposits in Arctic Canada representing the oldest taxonomically resolved red alga, implying primary plastid integration by at least 1.25 Ga, consistent with molecular clock estimates.^[92] Recent discoveries, such as the 2024 identification of the nitroplast—a nitrogen-fixing organelle derived from cyanobacterial endosymbiosis in marine algae—further illustrate symbiogenesis in modern eukaryotes.^[5]

Secondary Endosymbiosis

Process and Mechanisms

Secondary endosymbiosis begins with the phagocytic engulfment of a photosynthetic eukaryote, such as one bearing a primary plastid derived from an ancient cyanobacterium, by a non-photosynthetic heterotrophic eukaryotic host.^[93] This process, building on the precedent of primary endosymbiosis, integrates the endosymbiont's photosynthetic machinery into the host's cellular architecture through a series of reductive and integrative events.^[94] Following engulfment, the endosymbiont's plasma membrane fuses with the host's phagosomal membrane, resulting in complex plastids surrounded by three or four bounding membranes. The outermost membrane typically derives from the host's endomembrane system, while the inner two correspond to the primary plastid envelopes, and any additional membrane arises from the endosymbiont's original plasma membrane or further compartmentalization.^[95] These multilayered envelopes necessitate specialized protein import machineries, involving translocons that facilitate the transport of nuclear-encoded proteins across multiple barriers to maintain plastid function.^[96] In certain lineages retaining vestiges of the endosymbiont's autonomy, a nucleomorph forms as the reduced remnant of the engulfed eukaryote's nucleus, positioned between the second and third membranes of the complex plastid. This diminutive organelle, comprising a highly compacted genome, serves as an intermediate repository for genes essential to plastid maintenance before their relocation to the host nucleus. Nucleomorph genomes are characterized by extreme reduction, retaining only a few hundred genes organized into minimal chromosomes lacking introns and intergenic regions.^[97]^[98] Gene transfer plays a central role in the endosymbiont's integration, with extensive endosymbiotic gene transfer (EGT) relocating genetic material from the endosymbiont's nucleus—and often via the nucleomorph—to the host's nuclear genome. This process involves the physical escape of DNA fragments from the endosymbiont, their incorporation into host chromosomes, and the evolution of targeting signals on transferred genes to ensure protein reimport into the plastid. A second wave of EGT occurs, distinct from primary endosymbiosis, transferring genes originally from the primary plastid host to the secondary host nucleus, thereby consolidating control under the host genome.^[42]^[99]^[93] Metabolic integration ensues as the host gains access to photosynthesis-derived carbon fixation and electron transport, while the endosymbiont relinquishes independent metabolic pathways, becoming reliant on the host for nutrients and replication cues. This interdependence rewires cellular metabolism, with the plastid contributing not only to autotrophy but also to essential anabolic processes like fatty acid and amino acid synthesis, fully embedding it within the host's biochemical network. The loss of endosymbiont autonomy is marked by coordinated gene expression, where host nuclear factors regulate plastid division and function, ensuring synchronized cellular growth.^[100]^[40]^[101]

Examples in Eukaryotes

Secondary endosymbiosis refers to the process where a photosynthetic eukaryote, typically bearing a primary plastid derived from cyanobacteria, is engulfed by a non-photosynthetic eukaryote host, resulting in the integration of a secondary plastid surrounded by additional membranes. This event has occurred independently multiple times in eukaryotic evolution, leading to diverse lineages with complex plastids that often feature three or four bounding membranes. Key examples illustrate the variability in endosymbiotic integration, including the retention of vestigial endosymbiont nuclei called nucleomorphs in some cases, and the transfer of genes from the endosymbiont to the host nucleus. In cryptophytes, secondary endosymbiosis involved the engulfment of a red alga, producing plastids bounded by four membranes and retaining a nucleomorph—a reduced nucleus of the engulfed alga with a genome of approximately 500–700 kbp (e.g., 551 kbp in Guillardia theta) encoding around 30 plastid-targeted proteins.^[102] This nucleomorph resides between the outer two and inner two membranes, providing direct evidence of the endosymbiotic event. Cryptophytes, such as Guillardia theta, use these plastids for photosynthesis and phycobiliprotein-based light harvesting, with phylogenetic analyses confirming the red algal origin.^[103] Chlorarachniophytes represent an independent secondary endosymbiosis where a green alga was engulfed, yielding plastids with four membranes and a nucleomorph genome of about 373 kbp encoding roughly 17 plastid-targeted genes. Species like Bigelowiella natans exemplify this, with the nucleomorph located in the periplastid space, and molecular data supporting the green algal ancestry through shared pigment compositions and gene sequences. These plastids enable mixotrophic nutrition, combining photosynthesis with predation.^[104] Euglenids acquired their plastids through secondary endosymbiosis of a green alga, resulting in organelles bounded by three membranes without a nucleomorph, as the endosymbiont's nucleus was fully reduced. In organisms such as Euglena gracilis, these plastids store paramylon as a carbohydrate reserve and perform photosynthesis using chlorophylls a and b, with genomic evidence from nuclear genes of green algal origin confirming the event. This acquisition occurred independently from other green-secondary lineages. Dinoflagellates display remarkable diversity in secondary and higher-order endosymbioses, with many possessing peridinin-containing plastids derived from a secondary red algal endosymbiont, bounded by three membranes. For instance, peridinin dinoflagellates like Symbiodinium spp. integrate these plastids for autotrophy, often as endosymbionts in corals. Some dinoflagellates, such as Karenia brevis, have undergone tertiary endosymbiosis by engulfing haptophytes or other algae, leading to plastids with four membranes and further gene transfers. A few retain nucleomorph-like structures, underscoring ongoing endosymbiotic dynamics.^[105] Several chromalveolate groups, including haptophytes and ochrophytes (e.g., diatoms and brown algae), trace their plastids to a single ancient secondary endosymbiosis of a red alga, producing four-membrane plastids without nucleomorphs. In haptophytes like Emiliania huxleyi, these plastids support calcification and bloom formation in oceans, with extensive gene transfer from the endosymbiont evident in host nuclear genomes. Ochrophytes, comprising stramenopiles, similarly acquired red-derived plastids, enabling diverse ecological roles from phytoplankton to kelp forests. Phylogenetic reconstructions support this shared origin, with subsequent divergences.^[105] Apicomplexans, such as the malaria parasite Plasmodium falciparum, possess a non-photosynthetic secondary plastid called the apicoplast, derived from the engulfment of a red alga and bounded by four membranes, without a nucleomorph. The apicoplast plays essential roles in isoprenoid biosynthesis and fatty acid metabolism, making it a target for antimalarial drugs. Genomic evidence confirms its red algal origin through phylogenetic analysis of apicoplast-targeted proteins.^[106]

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