Fact-checked by Grok 2 weeks ago

B chromosome

B chromosomes, also known as supernumerary or accessory chromosomes, are dispensable genetic elements that occur in addition to the standard complement of essential chromosomes, termed A chromosomes, in the genomes of various eukaryotes.^[1] They are found in approximately 15% of eukaryotic species, spanning fungi, plants, and animals, with over 3,000 documented cases including well-studied examples in maize (Zea mays), rye (Secale cereale), and grasshoppers.^[2] ^[3] Unlike A chromosomes, B chromosomes typically do not undergo pairing or recombination during meiosis, exhibit irregular transmission often exceeding Mendelian ratios through mechanisms like meiotic drive, and vary in copy number from zero to dozens per cell across individuals in a population.^[1] ^[3] Structurally, B chromosomes are often smaller and predominantly heterochromatic, composed largely of repetitive DNA sequences such as satellite DNA, transposable elements, and ribosomal DNA (rDNA) arrays, though they may also incorporate euchromatic segments, pseudogenes, and even functional protein-coding genes derived from A chromosomes.^[2] ^[3] Their origins trace back to A chromosomes or other B chromosomes via processes like segmental duplication, transposition, or interspecific hybridization, leading to mosaicism where B chromosomes accumulate sequences from multiple genomic sources over evolutionary time.^[1] In some cases, they contain organelle-derived DNA or microRNAs that can influence host gene expression, and their gene content varies widely, with examples including histone genes in locusts or virulence factors in fungi.^[3] This structural diversity contributes to their persistence despite lacking essential functions, as they are non-vital for the organism's survival or reproduction.^[2] B chromosomes propagate through specialized accumulation mechanisms, primarily drive—a selfish transmission strategy that biases inheritance in their favor, such as nondisjunction during meiosis in maize or preferential segregation in pollen of grasses.^[1] ^[3] These can occur at mitotic, meiotic, or post-meiotic stages, sometimes at the cost of host fitness, leading to parasitic behavior where B chromosomes exploit the host's reproductive machinery.^[3] However, their effects on the host are multifaceted: while often neutral or deleterious—altering cell cycle, nutrient allocation, or fertility—they can confer adaptive advantages, such as enhanced stress resistance in plants or increased pathogenicity in fungi.^[2] In certain species, like cichlid fish, B chromosomes influence sex determination, highlighting their potential role in evolutionary innovation.^[3] From an evolutionary perspective, B chromosomes represent dynamic genomic parasites that may transition from selfish entities to integrated components, potentially evolving into essential chromosomes under relaxed selection pressures.^[1] High-throughput omics analyses have revealed their repetitive nature and gene activity, underscoring their impact on host genome stability and adaptation across taxa.^[2] Despite their dispensability, B chromosomes' persistence challenges traditional views of genome evolution, offering insights into genomic conflict and variability.^[3]

Overview

Definition and Characteristics

B chromosomes, also known as supernumerary or accessory chromosomes, are nonessential genetic elements that occur in addition to the standard complement of essential A chromosomes within the genome of certain eukaryotic organisms.^[1] These chromosomes are dispensable for normal viability and fertility, meaning individuals can survive and reproduce without them, and their presence varies widely among individuals and populations of the same species, ranging from zero to multiple copies.^[2] Unlike A chromosomes, which form the core set required for basic cellular and organismal functions, B chromosomes do not pair or recombine regularly with A chromosomes during meiosis.^[1] Key characteristics of B chromosomes include their frequent heterochromatic composition, which consists largely of repetitive DNA sequences, and their typically smaller size compared to A chromosomes, although this varies by species.^[2] They generally lack a complete set of genes necessary for independent viability, containing instead fragmented or partial gene sequences, though some may harbor functional elements related to their transmission.^[1] A hallmark feature is their ability to accumulate through non-Mendelian inheritance patterns, such as meiotic drive, which allows them to spread beyond expected segregation ratios despite providing no essential benefit to the host.^[2] B chromosomes are distinct from other chromosomal phenomena, such as marker chromosomes, which are abnormal structural variants derived directly from A chromosomes and often homologous to specific standard chromosomes, or trisomies, which involve extra copies of entire A chromosomes leading to gene dosage imbalances.^[1] In contrast, B chromosomes are not homologous to any A chromosome and represent truly supernumerary elements without disrupting the balanced A set.^[4] These properties have led to their identification in over 3,000 species across eukaryotes, spanning plants, animals, and fungi, highlighting their widespread but sporadic occurrence.^[5]

Prevalence Across Species

B chromosomes, as supernumerary elements distinct from the standard A chromosome complement, have been documented in approximately 15% of the eukaryotic species examined cytogenetically. This estimate, derived from early comprehensive reviews of karyotypic data, underscores their widespread but non-universal occurrence across fungi, plants, and animals. Recent compilations from specialized databases indicate that B chromosomes have been identified in over 2,800 species as of 2017, with updates extending to over 3,000 species as of 2024, spanning all major eukaryotic phyla.^[6] Among these, plants account for the majority of reports, comprising about 74% of documented cases, while animals represent roughly 26% and fungi less than 1%, highlighting a skewed distribution toward plant lineages. Patterns of B chromosome occurrence reveal variability both within and across populations. They are typically polymorphic, present in only a subset of individuals, with frequencies ranging from rare to over 50% in some populations, and individuals often carrying 0 to 15 copies.^[7] Their distribution tends to be geographically clustered, forming "hotspots" in certain regions or habitats rather than being evenly dispersed.^[7] Among flowering plants, B chromosomes are more prevalent in outcrossing species compared to self-incompatible or inbred ones, suggesting a link to mating systems that facilitate their transmission advantages.^[7] Early cytogenetic surveys from the 1960s to 1980s laid the foundation for understanding B chromosome distribution, identifying them in diverse taxa through microscopic analysis of chromosome spreads. Modern efforts, bolstered by databases such as B-chrom, have systematized this data up to the 2020s, enabling meta-analyses that account for sampling biases. Factors influencing prevalence include correlations with larger population sizes, which may buffer against deleterious effects, and associations with environmental stressors like pollution or climatic extremes, particularly in vertebrates.^[7]^[8] Additionally, meta-analyses have confirmed positive ties to genome size in plants and karyotype features like acrocentric chromosomes in mammals, indicating evolutionary and ecological drivers of their persistence.^[7]

Origin and Evolution

Evolutionary Origins

B chromosomes are hypothesized to originate as derivatives of standard A chromosomes through segmental duplications, translocations, or amplifications of heterochromatic regions, resulting in supernumerary elements that accumulate repetitive sequences over time.^[9] This derivation allows B chromosomes to initially carry functional gene fragments before evolving into largely non-essential, selfish genetic elements.^[10] Comparative genomic analyses provide strong evidence for this origin, showing B chromosomes as mosaics of A chromosome segments. For example, next-generation sequencing of the rye (Secale cereale) B chromosome revealed approximately 4,946 gene-derived sequences homologous to those on A chromosomes, particularly from rye chromosomes 3R and 7R, with blocks matching related grass genomes like barley.^[10] These findings indicate that the rye B arose from multiple recombination events and subsequent accumulation of repeats in pericentromeric areas.^[11] B chromosomes exhibit recurrent origins in diverse lineages, with no shared common ancestor across taxa, reflecting independent evolutionary events driven by genome instability.^[9] Recent studies suggest that chromoanagenesis—a process of massive chromosomal shattering and repair—may contribute to their formation following mitotic or meiotic segregation errors, leading to proto-B chromosomes.^[12] In plants, the rye B chromosome dates to 1.1–1.3 million years ago, aligning with the radiation of the Secale genus around 1.7 million years ago.^[10] In fish, such as species in the genus Astyanax, B chromosomes trace back to a common ancestor at least 4 million years ago, demonstrating long-term persistence through adaptive or neutral processes.^[13] In plants, B chromosomes frequently associate with polyploid events, where whole-genome duplications create genomic instability that promotes their formation via chromosome breakage and rearrangement.^[9] This link is evident in polyploid grasses like wheat and rye.^[14]

Mechanisms of Formation

B chromosomes arise through specific cytological and genetic processes that generate supernumerary elements from the standard A chromosome set, often involving instability and unequal segregation. One primary mechanism is nondisjunction during meiosis or mitosis, which leads to the initial accumulation of extra chromosomes by uneven distribution to daughter cells; for instance, in grasshoppers, mitotic instability results in nondisjunction, allowing B chromosomes to persist at low frequencies despite lacking essential genes.^[15] Another key process is the breakage-fusion-bridge (BFB) cycle, first described by Barbara McClintock in maize during the late 1930s and 1940s, where double-strand breaks in chromosomes lead to end-to-end fusions, forming dicentric structures that create anaphase bridges and subsequent breaks, generating fragmented derivatives that can stabilize as novel B chromosomes. This cycle has been implicated in the origin of B chromosomes. Translocation events further contribute to B chromosome formation by incorporating segments from A chromosomes, such as ribosomal DNA (rDNA) arrays or satellite repeats, into nascent B elements; in rye, for example, the B chromosome contains amplified rDNA sequences translocated from A chromosomes, enhancing its heterochromatic structure and stability.^[16] De novo B chromosome variants have been observed in laboratory settings, where cytomolecular analysis revealed origins from A chromosome duplications and rearrangements without prior B presence. The persistence of B chromosomes evolutionarily depends on a balance between formation and elimination mechanisms; for example, lagging B chromosomes during anaphase can form micronuclei, which are often excluded from the main nucleus and degraded, counteracting accumulation rates in species like Aegilops speltoides. This equilibrium ensures B chromosomes remain at low, variable frequencies across populations without overwhelming the A genome.

Structure and Composition

Morphology and Size

B chromosomes display a diverse array of morphologies, commonly appearing as telocentric or metacentric structures that distinguish them from the standard A chromosomes. They are typically smaller than A chromosomes, often comprising 10-50% of the length of the smallest A chromosomes, though this varies by species and population. For instance, in many grasshoppers, B chromosomes range from minute, dot-like elements to larger forms approaching the size of medium-sized autosomes.^[17]^[18]^[19] Size variation among B chromosomes is pronounced, spanning from tiny fragments visible only under high-resolution microscopy to nearly full-sized equivalents of A chromosomes in some cases. In the grasshopper Eyprepocnemis plorans, for example, standard B variants contain approximately 40-50% of the DNA content of the X chromosome, correlating with a relative length of about 20-50% depending on the variant. Similarly, in plants like maize (Zea mays), the B chromosome has a total length of 135 Mb, making it smaller than most A chromosomes, which range from 100-300 Mb, with a diminutive short arm and a long arm featuring distinct heterochromatic and euchromatic segments. In mammals such as the Siberian roe deer (Capreolus pygargus), B chromosomes are consistently smaller than the A set and often classified as microchromosomes.^[20]^[21] Cytologically, B chromosomes are heavily heterochromatic, resulting in dark staining with C-banding techniques that highlight pericentromeric and interstitial blocks of constitutive heterochromatin. This heterochromatin dominance contributes to their condensed appearance during mitosis. However, fluorescence in situ hybridization (FISH) reveals interspersed euchromatic regions in some B chromosomes, such as those containing repetitive DNA sequences homologous to A chromosomes or specific gene loci. These features are observed through standard karyotyping and light microscopy of metaphase spreads; for example, in metaphase preparations from Siberian roe deer cells, multiple small, heterochromatic B chromosomes (up to 14 per cell) are clearly distinguishable from the 70 A chromosomes. In plant species like maize, metaphase spreads similarly show the B chromosome's unique morphology, with its short arm and centromere appearing compact alongside the more extended A set.^[22]^[23]^[21]

Genetic and Molecular Composition

B chromosomes are primarily composed of repetitive DNA sequences, often constituting more than 80% of their total content, including satellite DNA, transposable elements, and tandem repeats.^[24] In many species, these repeats, such as LTR retrotransposons and DNA transposons, dominate the structure, with examples like the maize B chromosome featuring approximately 88.55% repeats, predominantly retrotransposons from the gypsy superfamily.^[25] Additionally, B chromosomes frequently retain ribosomal DNA (rDNA) clusters derived from A chromosomes, alongside other repetitive motifs like CentC and ZmBs near centromeric regions.^[26] While largely non-genic, they harbor a subset of functional genes transposed from standard chromosomes, including those involved in metabolism and cell cycle regulation, though these often exhibit reduced integrity and relaxed purifying selection.^[24] Genomic sequencing of B chromosomes remains challenging due to their repetitive nature and structural complexity, with full assemblies being rare until recent long-read technologies. Partial and chromosome-scale assemblies have provided key insights; for instance, the maize B chromosome was assembled to 135 Mb using PacBio HiFi and Hi-C, revealing no large syntenic regions with A chromosomes but confirming the presence of 1,124 protein-coding genes, of which 337 are expressed (FPKM >1) in leaf tissues.^[25] Similarly, in rye (Secale cereale), a 430 Mb pseudomolecule assembly from PacBio HiFi and Nanopore reads identified 1,292 transcriptionally active genes, including 799 protein-coding ones, amid a landscape dominated by satellite repeats like E3900 and D1100.^[27] These 2018–2025 studies across plants and animals, such as in fish (Astyanax species), highlight B chromosomes as derivatives of A chromosomes with extensive deletions and rearrangements, enriching our understanding of their molecular makeup, including unique centromeric satellite arrays like pZmBs in maize.^[24]^[25] Heterochromatin prevails in B chromosomes, particularly in pericentromeric regions enriched with repeats that contribute to meiotic stability and nondisjunction.^[28] In mammals like the Korean field mouse, B chromosomes add significant heterochromatin (up to 32% of the genome), featuring L1 elements and interstitial telomeres.^[28] However, euchromatic islands persist in some cases, containing active genes; for example, the rye B chromosome shows H3K4me3 marks on heterochromatic regions with functional genes, while maize Bs have distal euchromatic segments with metabolism-related loci.^[27]^[25] This mosaic composition underscores the retention of viable genetic material amid pervasive repeat accumulation. Post-2020 advances, driven by long-read sequencing rather than widespread CRISPR mapping, have unveiled novel mobile elements unique to B chromosomes, such as B-specific satellites in rye that influence drive mechanisms.^[27] In maize, these efforts confirmed the evolutionary divergence of B chromosomes over millions of years, with unique transposon proliferations not found in A sets.^[25] Such insights from high-impact studies emphasize the dynamic, repeat-heavy nature of B chromosomes while revealing pockets of functional euchromatin.^[24]

Biological Function

Parasitic and Drive Mechanisms

B chromosomes function as selfish genetic elements by employing meiotic drive to achieve non-Mendelian inheritance, transmitting to offspring at rates exceeding 50%, often through biased segregation that favors their inclusion in gametes.^[29] In some insect species, such as the parasitic wasp Nasonia vitripennis, this drive can approach 100% transmission via mechanisms that ensure the B chromosome is present in virtually all viable sperm. This selfish behavior allows B chromosomes to persist and accumulate in populations despite lacking essential genes for host viability.^[30] Several mechanisms underlie this drive, including nondisjunction during meiosis, where sister chromatids fail to separate properly, leading to both copies entering the same gamete. In female meiosis of certain grasshoppers like Eyprepocnemis plorans, nondisjunction directs B chromosomes preferentially toward the egg nucleus.^[31] Preferential segregation is another common strategy, observed in grasshoppers such as Myrmeleotettix maculatus, where univalent B chromosomes migrate toward the pole forming the functional gamete more frequently than expected by chance.^[30] Additionally, some B chromosomes induce the killing or elimination of gametes lacking them; for instance, in Nasonia vitripennis, the paternal sex ratio (PSR) B chromosome disrupts the paternal genome in zygotes without PSR, effectively eliminating competing genetic material and securing its transmission. The parasitic nature of B chromosomes manifests through fitness costs to the host upon accumulation, including reduced fertility and vigor. In plants like rye (Secale cereale), high numbers of B chromosomes (e.g., more than two) progressively decrease pollen fertility and seed set, as the extra chromatin burdens cellular processes.^[30] Similarly, in maize (Zea mays), individuals with multiple B chromosomes exhibit impaired vigor and fertility, counterbalancing the drive advantage.^[29] Population genetics models describe the equilibrium frequency of B chromosomes as a balance between their transmission advantage (drive) and elimination rates due to fitness costs from selection or irregular transmission. A seminal model for preferential segregation in Lilium callosum analyzes this balance, ensuring B chromosomes neither fix nor vanish entirely in populations, maintaining polymorphism.^[32]^[30]

Adaptive and Beneficial Roles

While B chromosomes are often viewed as selfish genetic elements, emerging evidence suggests they can provide adaptive advantages to host organisms, particularly under specific environmental pressures. One hypothesized benefit is their role in buffering genetic variation, where B chromosomes neutralize variability induced by heterogeneous conditions, such as differing soil types, thereby stabilizing phenotypic outcomes in plants like Trigonella foenum-graecum.^[33] Additionally, B chromosomes have been linked to enhanced stress resistance; for instance, in the plant Allium schoenoprasum, seeds carrying B chromosomes exhibit superior germination rates under drought conditions compared to those without, potentially through gene dosage effects that amplify stress-response pathways.^[34] In rye (Secale cereale), the presence of B chromosomes upregulates heat shock proteins like tE3900 by up to 40-fold during male sporogenesis under thermal stress, conferring greater tolerance.^[35] Specific functional contributions of B chromosomes further support their beneficial roles. In some species, B chromosomes amplify ribosomal DNA (rDNA) arrays, enabling nucleolar organization and rRNA transcription essential for cellular function; in the grasshopper Eyprepocnemis plorans, B-chromosome rDNA is actively transcribed, forming functional nucleoli in a dosage-dependent manner, with activity observed in up to 100% of individuals carrying three B chromosomes.^[36] Moreover, certain B chromosomes harbor active genes that provide direct advantages, such as in fungi where dispensable chromosomes carry cytochrome P-450 genes (pda) enabling detoxification of phytoalexins like pisatin, thus enhancing antibiotic and toxin resistance in pathogens like Nectria haematococca.^[37] Recent genomic analyses confirm that such functional genes persist on B chromosomes across fungal lineages, contributing to adaptive pathogenicity and environmental resilience.^[3] Experimental evidence underscores these benefits, particularly when B chromosomes are absent or removed. In field and lab studies, individuals lacking B chromosomes show reduced fitness under stress; for example, in Avena sativa (oats), small accessory fragment chromosomes carrying resistance genes (Pc-15) enhance survival against rust pathogens (Puccinia coronata f. sp. avenae), though these differ from typical B chromosomes.^[38]^[39] A 2022 analysis of rye B chromosomes revealed that low B copy numbers (1-2) optimize gene expression for stress adaptation without incurring costs, and artificial or natural elimination in root tissues leads to impaired development under variable conditions.^[40] Recent studies as of 2024-2025, including high-resolution sequencing, continue to uncover functional genes on B chromosomes that modulate host responses to environmental stresses across taxa.^[41] These findings highlight how B chromosome removal can exacerbate vulnerability in challenging habitats. Evolutionarily, B chromosomes may transition from parasitic entities to symbiotic components, especially when present in low numbers that balance transmission drive with host benefits. In lineages like cichlid fish (Astatotilapia latifasciata), B chromosomes have integrated functional roles, evolving into essential elements that support adaptation without overwhelming the genome.^[42] This shift is evident in their conservation of adaptive genes over time, suggesting that 1-2 B copies represent an optimal state for long-term persistence and mutualistic interactions.^[43]

Occurrence in Organisms

In Plants

B chromosomes are particularly prevalent among plant species in the Poaceae (grasses) and Liliaceae (lilies) families, where they occur in a notable proportion of taxa. Across plants overall, they have been documented in over 2,500 species, representing a notable proportion of those with known chromosome numbers (as of 2023).^[6] In grasses such as Aegilops speltoides, individuals can harbor up to eight B chromosomes alongside the standard 14 A chromosomes, with their frequency often increasing in polyploid lineages. This variation with ploidy level is evident in polyploid grasses like wheat (Triticum aestivum), where B chromosomes can accumulate to 20 copies without disrupting basic viability at low numbers. Recent genomic studies as of 2024 have further documented B chromosomes in polyploid wheat, enhancing understanding of their occurrence in crop species.^[44]^[45]^[46]^[44] Prominent examples of B chromosomes in plants include those in rye (Secale cereale), a diploid grass, where up to six B chromosomes influence meiotic behavior by altering A chromosome pairing and nondisjunction, often leading to reduced pollen fertility. These effects in rye extend to agronomic traits, with higher B chromosome numbers correlating with decreased seed set and grain yield due to impaired vigor and fertility. Similarly, in maize (Zea mays), another grass, B chromosomes—capable of reaching up to 34 copies—impact kernel development through instability during early endosperm formation, resulting in variegated or small-kernel phenotypes when present in elevated numbers. In lilies (Lilium spp.), B chromosomes exhibit meiotic drive, preferentially segregating to gametes and maintaining population frequencies despite lacking essential genes.^[45]^[47]^[26]^[48] B chromosomes in plants frequently alter host phenotypes, particularly at higher copy numbers, by exerting parasitic effects that reduce overall fitness. For instance, they can diminish seed set and fertility in grasses like Dactylis glomerata, where one to 12 B chromosomes progressively impair reproductive output. Phenotypic changes may also include modifications to floral traits, such as reduced flower size in certain outcrossing species, linked to increased genome size and heterochromatin load. Recent studies have highlighted potential beneficial roles under stress; in rye, B chromosomes enhance heat tolerance during male sporogenesis, protecting meiocytes from elevated temperatures.^[49]^[50]^[51] Post-2020 epigenomic research has addressed longstanding gaps in understanding B chromosome regulation, particularly in polyploids. In rye, B chromosomes display distinct histone modifications, including H3K4me3 and H3K27me3, which contribute to localized gene silencing and influence A chromosome expression via small RNA pathways. In polyploid wheat, B chromosomes undergo gene erosion and epigenetic silencing, with repetitive elements methylated to prevent interference with A genome stability, as revealed by chromatin mapping and transcriptomic analyses. These findings underscore how epigenetic mechanisms, such as DNA methylation and histone variants, facilitate B chromosome persistence in complex polyploid genomes without major deleterious impacts at low frequencies.^[52]^[40]^[44]^[53]

In Animals

B chromosomes are widespread in animal taxa, with reports documenting their presence in over 700 species across insects, mammals, and fishes.^[54] They occur frequently in insects, particularly within the order Orthoptera where approximately 10% of studied species harbor them, showing non-uniform distribution among superfamilies with hotspots in certain lineages.^[55] In fish, B chromosomes are notably prevalent in cichlids, where they often derive from sex chromosome material and appear in up to 85% of individuals in some populations, typically numbering from one to eight per cell.^[56]^[57] Prominent examples illustrate their distribution and impacts in animals. In the grasshopper Myrmeleotettix maculatus (Orthoptera: Acrididae), B chromosomes exhibit clinal variation in frequency across populations, with higher prevalence linked to more favorable environmental conditions such as warmer, drier habitats, suggesting a role in local adaptation.^[58]^[59] In mammals, the Siberian roe deer (Capreolus pygargus) displays up to 14 B chromosomes that are readily visible in karyotypes, often appearing as small, heterochromatic elements distinct from the standard 70-chromosome complement.^[60] These Bs in roe deer incorporate pseudogenized sequences and show independent evolutionary origins from those in related cervids.^[61] B chromosomes in animals frequently influence reproduction, often through drive mechanisms that distort inheritance patterns. In cichlid fishes, such as those in Lake Malawi, Bs cause female-biased sex ratios by preferentially accumulating in female offspring, altering sex determination via functional genes on the chromosome.^[56] High numbers of Bs generally reduce fertility, as seen in Lake Malawi cichlids where increased B copy numbers correlate with decreased reproductive success, though effects are minimal at low frequencies.^[62] Recent studies from 2021 to 2024 have utilized Drosophila melanogaster models to explore B chromosome drive, revealing a female meiotic drive suppression system where non-driving Bs interact with genetic backgrounds to restore balanced transmission, providing insights into evolutionary suppression of selfish elements.^[63]^[64] In certain animal groups, B chromosomes exhibit unique associations with ecological and applied contexts. They appear in social insects like hymenopterans, though less frequently documented than in solitary species, potentially influencing colony-level genetics in ants and bees.^[65] In aquaculture-relevant species such as cichlids, Bs' drive properties raise implications for breeding programs, where uncontrolled transmission could affect stock management.^[62] Furthermore, the selfish drive of Bs in insects has sparked interest in their potential adaptation for pest control strategies, analogous to gene drive technologies that could suppress invasive populations by biasing inheritance against fitness.^[66]

In Fungi

B chromosomes, also known as supernumerary or accessory chromosomes, exhibit widespread presence-absence polymorphisms in fungal species, particularly within the Ascomycetes phylum, where they contribute to intraspecific genome variation. These dispensable elements are non-essential for basic cellular functions but can influence phenotypic traits in specific environments. For instance, the wheat pathogen Zymoseptoria tritici harbors up to eight small dispensable chromosomes, ranging from 0.39 to 0.77 Mb, which display high variability across strains and are absent in some isolates without affecting saprophytic growth.^[67]^[68]^[69] In plant-pathogenic fungi, B chromosomes often carry genes that enhance host interaction and survival. A prominent example is Haematonectria haematococca (synonym Nectria haematococca), where supernumerary chromosomes encode resistance to pea-derived antimicrobial compounds, such as pisatin, enabling the fungus to detoxify host defenses and promote infection. Similarly, in Zymoseptoria tritici, accessory chromosomes harbor effector genes that modulate virulence, with their presence correlating to increased aggressiveness on wheat hosts. These elements typically feature higher repeat content, gene duplications, and lower GC bias compared to core chromosomes, positioning them as dynamic repositories for adaptive alleles.^[70]^[71]^[72] Functional studies indicate that fungal B chromosomes can boost pathogenicity by facilitating effector deployment or toxin resistance, while also aiding nutrient acquisition in nutrient-limited host tissues through specialized metabolic genes. Recent genomic sequencing efforts, including analyses from 2022, have illuminated their role as gene reservoirs, revealing elevated transposable element densities and duplicated loci that support rapid evolution of virulence factors without disrupting essential genome stability. For example, in Magnaporthe oryzae, mini-chromosomes serve as hotspots for effector genes, enhancing adaptation to rice hosts.^[73]^[5]^[74] Research on fungal B chromosomes began with cytogenetic approaches in the early 2000s, which mapped their transmission and structural features using pulsed-field gel electrophoresis to identify dispensable elements in pathogens like Nectria haematococca. These foundational studies highlighted non-Mendelian inheritance and polymorphisms but left gaps in gene content and function. Post-2020 functional genomics, leveraging long-read sequencing and CRISPR-based knockouts, has addressed these by confirming dispensability through loss-of-function assays and revealing epigenetic regulation, such as H3K27me3 enrichment, that silences B chromosome genes under non-host conditions. This shift has underscored their conditional benefits in pathogenicity without essential roles in axenic growth.^[75]^[76]^[77]^[78]

References

[1]
What is a B chromosome? Early definitions revisited - PMC
Apr 16, 2024 · B chromosomes (Bs) are enigmatic heritable elements that have been identified in many fungal, plant, and animal species (Gutiérrez et al. 2023).Missing: review | Show results with:review
[2]
The Modern View of B Chromosomes Under the Impact of High ...
Feb 13, 2019 · B chromosomes (Bs): Chromosomes that are not essential parts of the genome and therefore not present in all individuals in a population of a ...
[3]
Comprehending the dynamism of B chromosomes in their journey ...
Represented singly or in multiple copies, B chromosomes are largely heterochromatic but also contain euchromatic and organellar segments. Although B chromosomes ...Missing: paper | Show results with:paper
[4]
Human supernumeraries: are they B chromosomes?
Aug 18, 2004 · Most human supernumerary chromosomes (HSCs) derive from the A chromosome set, but also the specific A chromosome from which most of them arose.
[5]
The distribution of B chromosomes across species - PubMed
In this review we look at the broad picture of how B chromosomes are distributed across a wide range of species. We review recent studies of the factors ...
[6]
The genetic mechanism of B chromosome drive in rye illuminated by ...
Nov 8, 2024 · As of 2023, B chromosomes have been discovered in 2951 species from all eukaryotic phyla. Most B chromosomes confer no detectable selective ...
[7]
The distribution of B chromosomes across species - Karger Publishers
Aug 18, 2004 · Abstract. In this review we look at the broad picture of how B chromosomes are distributed across a wide range of species.
[8]
Origin, Behaviour, and Transmission of B Chromosome with Special ...
Feb 18, 2019 · It has been estimated that Bs occur in 15% of eukaryotic species [2] and most of these are plants [14]. As far as animals are concerned, Bs are ...
[9]
Evolution and biology of supernumerary B chromosomes - PMC - NIH
Depending on the species, the half-life of an active duplicated gene becoming mutated or lost was estimated to be 2–7 million years [86]. Considering this ...
[10]
Selfish supernumerary chromosome reveals its origin as a ... - PNAS
Supernumerary B chromosomes are optional additions to the basic set of A chromosomes, and occur in all eukaryotic groups. They differ from the basic ...Missing: across | Show results with:across
[11]
High‐copy sequences reveal distinct evolution of the rye B ...
Apr 24, 2013 · We proposed that rye Bs are descended from rearrangements of chromosomes 3RS and 7R, with subsequent accumulation of repeats and genic fragments ...Introduction · Materials and Methods · Results · Discussion
[12]
Long-term persistence of supernumerary B chromosomes in multiple ...
Mar 19, 2021 · ... B chromosomes reported in several fish species of the genus Astyanax arose in a common ancestor at least 4 million years ago. We generated ...
[13]
B‐chrom: a database on B‐chromosomes of plants, animals and fungi
Jul 25, 2017 · We present here B-chrom (www.bchrom.csic.es), an online database with comprehensive information on B chromosomes (Bs) for plants, animals ...
[14]
https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.14723
[15]
https://doi.org/10.1007/BF00326786
[16]
Current knowledge on B chromosomes in natural populations of ...
Aug 18, 2004 · B chromosomes showed a diverse morphology, including telocentric to metacentric structure. There is no detailed banding or ultrastructural ...
[17]
B Chromosomes in Grasshoppers: Different Origins and Pathways to ...
Origin of Dot-Like B. The Bs in grasshoppers vary in size from minute dot-like elements to a large chromosome equal to the largest autosome (Figure 1, Figure 2 ...Missing: variation | Show results with:variation
[18]
Evolution of B Chromosomes: From Dispensable Parasitic ... - Frontiers
... B chromosomes occur in odd numbers (Camacho et al., 2004). B chromosomes show high variability in their size across species. They can be similar in size to ...Abstract · Introduction · Conclusion
[19]
DNA Amount of X and B Chromosomes in the Grasshoppers ...
Mar 8, 2011 · We analyzed the DNA amount in X and B chromosomes of 2 XX/X0 grasshopper species (Eyprepocnemis plorans and Locusta migratoria), by means of ...<|control11|><|separator|>
[20]
Genome assembly of the maize B chromosome provides insight into ...
Mar 6, 2025 · B chromosomes contribute to the genetic variation in numerous eukaryotes. Yet their genetic and epigenetic characteristics, as well as their ...Missing: paper | Show results with:paper
[21]
B Chromosomes in Populations of Mammals Revisited - PMC - NIH
Oct 9, 2018 · In this genus, Bs appear in five out of eight species [51]. For instance, in M. americana, Abril et al. [127] found Bs in all 18 studied animals ...
[22]
Genes on B chromosomes of vertebrates - Molecular Cytogenetics
Dec 17, 2014 · Here we review recent findings of gene-carrying genomic segments on B chromosomes from different vertebrate groups.
[23]
Comparative FISH Analysis of Distribution of B Chromosome ...
In the B chromosomes, C-positive regions contained homologous DNA repeats present also in some C-positive A chromosome regions. Most C-negative regions ...
[24]
B chromosomes of multiple species have intense evolutionary ...
Sep 23, 2020 · This study reveals the genomic structure, composition and function of Bs, which provide new insights for theories of B chromosome evolution.
[25]
Sequence of the supernumerary B chromosome of maize ... - PNAS
B chromosomes are nonvital chromosomes found in thousands of plants and animals that persist through various drive mechanisms. The drive mechanism of the ...
[26]
The genetic mechanism of B chromosome drive in rye illuminated by ...
Nov 8, 2024 · The B chromosome of rye – a model system for nearly a century – undergoes targeted nondisjunction during first pollen mitosis, favouring ...Missing: irradiation | Show results with:irradiation
[27]
Sequence Composition and Evolution of Mammalian B Chromosomes
Oct 10, 2018 · Here, we consider mammalian Bs, taking into account data on their DNA sequencing, transcriptional activity, positions in nuclei of somatic and meiotic cells,
[28]
https://www.mdpi.com/2073-4425/9/10/490
[29]
https://doi.org/10.3389/fpls.2017.00210
[30]
https://doi.org/10.3390/genes9080388
[31]
https://doi.org/10.3390/genes9100509
[32]
Buffering effect of B-chromosome system of Trigonella foenum ...
The conclusion that can be drawn from this experiment is that B-chromosomes act as a buffering agent, neutralising the variability effect of different soil ...Missing: variation | Show results with:variation
[33]
https://link.springer.com/article/10.1007/BF00265609
[34]
https://doi.org/10.1038/hdy.1989.78
[35]
B-Chromosome Ribosomal DNA Is Functional in the Grasshopper ...
May 3, 2012 · Therefore, B-chromosome rDNA in E. plorans is functional since it is actively transcribed to form the nucleolus attached to the B chromosome.
[36]
A Fungal Gene for Antibiotic Resistance on a Dispensable - Science
... functional genes can be found on B chromosomes. Unstable B chromosomes may be one mechanism for generating pathogenic variation in fungi. Formats available.
[37]
https://www.science.org/doi/10.1126/science.1763326
[38]
Rye B chromosomes differently influence the expression of A ...
Jul 4, 2022 · According to earlier research, reductions in plant fitness are pronounced when the B chromosome mass exceeds 20% of DNA in the nucleus.Rye B Chromosomes... · Material And Methods · Go Enrichment Analyses...<|separator|>
[39]
https://doi.org/10.2135/cropsci1973.0011183X001300060002x
[40]
Evolution of B Chromosomes: From Dispensable Parasitic ... - PMC
Dec 9, 2021 · Based on the number of species studied so far it has been estimated that approximately 15% of eukaryotic species have B chromosomes (Beukeboom, ...
[41]
The non-Mendelian behavior of plant B chromosomes - PMC - NIH
Apr 12, 2022 · So far, B chromosomes have been found in 2087 plant species, accounting for 2.68% of 77,958 species with a known chromosome number (D'Ambrosio ...Missing: prevalence | Show results with:prevalence
[42]
B chromosome first—mechanisms behind the drive of B ... - Phys.org
Jun 21, 2019 · Ae. speltoides is known to carry up to 8 B chromosomes in addition to its 14 standard A chromosomes. Using comparative genomics, the scientists ...
[43]
B Chromosomes in Wheat: Evolution, Functions and Breeding ...
They have a significant impact on seed germination and pollen fertility in wheat. According to earlier research, there are differences in fertility between 2B ...
[44]
Rye (Secale cereale) supernumerary (B) chromosomes associated ...
Nov 5, 2016 · Relevantly, these extra chromosomes have various effects on mitotic and meiotic A chromosome behaviour. For example, Bs alter rDNA ...
[45]
Instability of the maize B chromosome - SpringerLink
Much less commonly the chromosome is unstable during early development of the kernel. Instability in the kernel produces recessive sectors in either the ...
[46]
The effects of B-chromosomes on vigour and fertility in Dactylis hybrids
B-chromosomes ranging in number from one to 12 had a deleterious effect on the vigour and fertility of a population of tetraploid hybrids between:.
[47]
B chromosomes and genome size in flowering plants - PubMed
Here we show that the presence of B chromosomes correlates to and varies strongly and positively with total genome size (excluding the Bs and corrected for ...Missing: seed set
[48]
Rye (Secale cereale) supernumerary (B) chromosomes associated ...
Nov 5, 2016 · Taken together, these results are the first indication that rye B chromosomes have implications on heat tolerance and may protect meiocytes ...
[49]
https://www.tandfonline.com/doi/pdf/10.1080/0028825X.1968.10428579
[50]
The bread wheat epigenomic map reveals distinct chromatin ...
Jul 15, 2019 · We herein present a systematic epigenomic map for the annotation of cis-regulatory elements in the bread wheat genome, which provides new insights.
[51]
B chromosomes of multiple species have intense evolutionary ...
Sep 23, 2020 · B chromosomes have been reported in 744 animal species including insects, mammals and fishes (http://www.bchrom.csic.es [5]). In addition to ...
[52]
Prevalence of B chromosomes in Orthoptera is associated ... - PubMed
B chromosomes have been reported in 191 of these species. Bs are not uniformly distributed among orthopteran superfamilies, with evident hotspots in the ...Missing: fish cichlids
[53]
B Chromosomes Have a Functional Effect on Female Sex ...
B chromosomes, which are present in addition to standard chromosomes, were found at a high prevalence rate (85%) in these cichlids.Missing: insects | Show results with:insects<|control11|><|separator|>
[54]
The B chromosomes of the African cichlid fish Haplochromis ...
Jan 5, 2010 · In general, the B chromosomes of fishes vary in number from one to eight [13–15]. The size of the B chromosomes can vary greatly, from macro- ...
[55]
RAPID MOVEMENT OF A B-CHROMOSOME FREQUENCY CLINE ...
The ecological genetics of the polymorphism for B-chromosomes in the grasshopper Myrmeleotettix maculatus (Thunb.) (Orthoptera: Acrididae). Ph.D. thesis ...
[56]
[PDF] Macrogeographic patterns in B-chromosome and inversion ... - HAL
May 11, 2020 · hautes fréquences du chromosome B dans les populations naturelles sont associées à des conditions plus favorables de milieu. orthoptère / ...
[57]
Transcription of a protein-coding gene on B chromosomes of the ...
Aug 6, 2013 · The only derived karyotypic character of the Siberian roe deer is the presence of B chromosomes [23], the number of supernumerary elements ...
[58]
Contrasting origin of B chromosomes in two cervids (Siberian roe ...
Aug 11, 2016 · The B chromosomes of the Siberian roe deer incorporate only two regions less than 2-Mbp in size characterized by a high pseudogenisation and ...
[59]
B chromosomes of Lake Malawi cichlid fish | Scientific Reports
Dec 27, 2019 · Fertility typically decreases with the presence of B chromosomes, but this and other effects on fitness are not usually significant unless an ...
[60]
B chromosomes reveal a female meiotic drive suppression system in ...
Jun 5, 2023 · Selfish genetic elements use a myriad of mechanisms to drive their inheritance and ensure their survival into the next generation, often at ...
[61]
Scientists discover the dynamics of an 'extra' chromosome in fruit flies
May 4, 2023 · B chromosomes reveal a female meiotic drive suppression system in Drosophila melanogaster. Current Biology, 2023; DOI: 10.1016/j.cub.2023.04 ...
[62]
Chromosome study of the Hymenoptera (Insecta)
Nov 1, 2023 · ... B chromosomes hybridizes only with these elements. ... The Honeybee Genome Sequencing Consortium (2006) Insights into social insects from the ...
[63]
The value of existing regulatory frameworks for the environmental ...
... B chromosomes, homing ... In this section, we briefly review two broad classes of synthetic gene drives and their potential for insect pest control.Review · Introduction · Synthetic Gene Drive...
[64]
Cell biology of Zymoseptoria tritici: Pathogen cell organization ... - NIH
Z. tritici carries 21 chromosomes (Goodwin et al., 2011). Eight of the smallest chromosomes appear to be non-essential (dispensable chromosomes) and are ...
[65]
Histone modifications rather than the novel regional centromeres of ...
Oct 1, 2015 · In fungi, these “accessory” or “dispensable” chromosomes are present at different frequencies in populations and are usually characterized by ...
[66]
Overview of genomic and bioinformatic resources for Zymoseptoria ...
... dispensable chromosomes ranging from 0.39 to 0.77 Mb – were not required for saprophytic growth i.e. were dispensable or accessory chromosomes (ACs) ...
[67]
The Genome of Nectria haematococca - NIH
Aug 28, 2009 · Subsequent research has demonstrated that in N. haematococca MPVI, genes on these chromosomes are involved in resistance to plant antimicrobials ...
[68]
(PDF) The Genome of Nectria haematococca - ResearchGate
These supernumerary chromosomes contain more repeat sequences, are enriched in unique and duplicated genes, and have a lower G+C content in comparison to the ...
[69]
Forward Genetics Approach Reveals Host Genotype-Dependent ...
Nov 28, 2017 · The fungal wheat pathogen Zymoseptoria tritici possesses a large complement of accessory chromosomes showing presence/absence polymorphism ...
[70]
Variability of chromosome structure in pathogenic fungi — of 'ends ...
Some fungal B chromosomes are 'pathogenicity' chromosomes. Subtelomeric regions and B chromosomes share widespread H3K27me3.Missing: prevalence | Show results with:prevalence
[71]
Comprehending the dynamism of B chromosomes in their journey ...
Represented singly or in multiple copies, B chromosomes are largely heterochromatic but also contain euchromatic and organellar segments. Although B chromosomes ...
[72]
Rapid mini‐chromosome divergence among fungal isolates causing ...
Nov 20, 2023 · Compared with core chromosomes, supernumerary mini-chromosomes in M. oryzae have a lower gene density and higher levels of repetitive sequences.
[73]
B-chromosome evolution | Philosophical Transactions of the Royal ...
B chromosomes are extra chromosomes to the standard complement that occur in many organisms. They can originate in a number of ways including derivation ...
[74]
Full article: Cytological karyotyping and characterization of a 410 kb ...
In this study we present a reliable karyotype of this important plant pathogen, N. haematococca MPI, by combining PFGE and GTBM. In addition, a mini-chromosome ...
[75]
Non-Mendelian transmission of accessory chromosomes in fungi
Jul 26, 2022 · In this review, we provide an overview of the possible mechanisms that can cause the non-Mendelian transmission or meiotic drives of fungal accessory ...
[76]
The Fusarium verticillioides minichromosome negatively regulates ...
Aug 2, 2025 · The minichromosome negatively modulates fumonisin biosynthesis during maize infection, and its deletion increases fumonisin levels.<|separator|>