Fact-checked by Grok 2 weeks ago

Reproductive isolation

Reproductive isolation refers to the genetically based mechanisms that reduce or prevent between populations or , thereby maintaining their genetic distinctiveness and serving as a cornerstone of in . These barriers can be broadly classified into prezygotic mechanisms, which prevent mating or fertilization, and postzygotic mechanisms, which reduce the viability or fertility of hybrid offspring. At its core, reproductive isolation underpins the biological species concept, first articulated by , which defines a as a group of actually or potentially interbreeding natural populations that are reproductively isolated from other such groups. Prezygotic barriers include temporal isolation, where species breed at different times; behavioral isolation, involving incompatible mating rituals; mechanical isolation, due to mismatched genitalia or pollinators; habitat isolation, where populations occupy different environments; and gametic isolation, where sperm and egg are incompatible. Postzygotic barriers encompass hybrid inviability, where embryos fail to develop; hybrid sterility, as seen in mules from horse-donkey crosses; and hybrid breakdown, where later generations are unfit. These mechanisms can evolve gradually through , , or , often reinforced in sympatric or allopatric contexts to prevent costly hybridization. The study of reproductive isolation has profound implications for understanding , as it quantifies how genetic differences impede and facilitate the formation of new , with applications in fields from to . For instance, in , can instantly create postzygotic isolation, leading to rapid events. Overall, reproductive isolation not only delineates boundaries but also highlights the dynamic processes driving .

Definition and Overview

Reproductive isolation encompasses the biological mechanisms that prevent or limit interbreeding between distinct populations or , thereby preserving their genetic integrity and facilitating evolutionary . These mechanisms are categorized into two primary types: prezygotic barriers, which act prior to fertilization by hindering attempts or the union of gametes, and postzygotic barriers, which operate after fertilization by diminishing the survival, development, or of hybrid offspring. This underscores the multifaceted nature of isolation as a quantitative process influenced by genetic, ecological, and behavioral factors. The foundational development of reproductive isolation as a central in traces back to the mid-20th century, building on earlier genetic insights. laid early groundwork in the 1930s through experiments on fruit flies, where he identified genetic bases for hybrid sterility and other isolating factors, culminating in his 1937 book Genetics and the Origin of Species, which introduced "isolating mechanisms" as genotypic differences reducing between populations. advanced this framework during the Modern Synthesis in the 1940s, particularly in his 1942 book Systematics and the Origin of Species, where he positioned reproductive isolation as the defining criterion of the biological concept, describing as "groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups." Reproductive isolation plays a pivotal role in by enforcing genetic boundaries that allow populations to evolve independently, thereby contributing to the generation and maintenance of . It provides a critical lens for delineating limits in systems. Modern genomic analyses since 2020 have illuminated its nuances, revealing that boundaries are often porous due to intermittent and , which can promote adaptive evolution while isolation mechanisms ultimately prevent genetic homogenization. For instance, studies in demonstrate widespread across lineages, highlighting how incomplete isolation drives diversification amid ongoing exchange.

Prezygotic Barriers

Habitat Isolation

Habitat isolation occurs when populations of closely related occupy different ecological niches or physical locations, thereby preventing encounters between potential mates and reducing interbreeding opportunities. This prezygotic barrier arises from physical separation due to distinct habitats, such as versus terrestrial environments, or finer-scale differences like microhabitats within the same geographic area. For instance, may diverge by preferring specific substrates, water depths, or vegetation types that limit overlap in their ranges. The mechanisms underlying habitat isolation can originate through allopatric processes, where geographic barriers like , mountains, or divides physically separate populations, leading to independent evolution of habitat preferences. In sympatric scenarios, isolation emerges from niche divergence driven by ecological selection, where populations adapt to different resources or conditions within the same area, such as varying food sources or environmental gradients. This divergence often begins as an to local conditions but incidentally promotes reproductive isolation by minimizing contact. A classic example is the fly (Rhagoletis pomonella), where ancestral hawthorn-infesting populations shifted to apple hosts following the introduction of apple trees to around 200 years ago, resulting in sympatric races that prefer different host plants and exhibit reduced mating due to host-specific odor preferences and aggregation. Similarly, in African rift lakes, fishes like those in have diverged by depth preferences, with deep-water species (Diplotaxodon spp.) isolated in benthic zones featuring limited light spectra, leading to genetic differentiation and despite . Habitat isolation frequently serves as an initial step in , as ecological selection on habitat preferences generates barriers that accumulate over time and reinforce divergence. Studies of hybrid zones, such as those in East African cichlids, demonstrate how shifts in habitat use along depth gradients maintain isolation even in secondary contact, with of traits enhancing prezygotic barriers. Recent analyses (2020–2025) highlight that such habitat-driven isolation strengthens linearly with , underscoring its role in ongoing processes.

Temporal Isolation

Temporal isolation is a prezygotic reproductive barrier in which differences in the timing of critical reproductive events, such as seasons, flowering periods in , or daily activity cycles, reduce or prevent the overlap in availability between or populations, thereby limiting interbreeding. This mechanism arises when populations diverge in their phenological responses, ensuring that individuals are reproductively active at non-overlapping times despite occupying the same geographic area. The primary drivers of temporal isolation are environmental cues, including temperature fluctuations and photoperiod (day length), which synchronize reproductive timing with seasonal conditions optimal for offspring survival. These cues trigger physiological changes, such as gonadal maturation in animals or floral induction in plants, leading to shifts in reproductive . Under divergent selection pressures, such as varying local climates, temporal isolation can evolve rapidly; recent meta-analyses indicate that in response to these cues accelerates the development of reproductive barriers during early stages of . Illustrative examples include the northern red-legged frog (Rana aurora), which breeds earlier in the season than the foothill yellow-legged frog (Rana boylii), minimizing hybridization opportunities despite shared habitats. In periodical cicadas of the genus Magicicada, species with 13-year life cycles (M. tredecim) and 17-year cycles (M. septendecim) emerge asynchronously, preventing mating despite overlapping habitats and serving as a strong temporal barrier that contributes to speciation. Among plants, three sympatric orchid species in tropical rainforests (Encyclia, Epidendrum, and Prosthechea) bloom at distinct times within the same season—early, mid, and late—driven by subtle differences in photoperiod sensitivity, which isolates pollen transfer. Temporal isolation is quantified using overlap indices from phenological data, such as the proportion of shared reproductive activity periods or models estimating pollen flow reduction based on flowering time differences; for instance, in sunflowers, a 10-day shift in bloom timing can decrease interpopulation by over 90%. This barrier plays a key role in by enabling divergence without geographic separation, as seen in stonefly populations (Leuctra hippopus) where seasonal emergence shifts create complete reproductive isolation within shared streams. Such temporal mismatches can compound with differences to further reinforce isolation ecologically.

Behavioral Isolation

Behavioral isolation is a prezygotic reproductive barrier that arises from differences in mating behaviors, such as displays, pheromones, or vocalizations, which prevent individuals from different from recognizing each other as potential mates. These behavioral divergences ensure that attempts are typically directed toward conspecifics, thereby reducing interspecific hybridization without requiring physical or temporal separation. Several mechanisms underlie the evolution of behavioral isolation. The sensory drive hypothesis posits that environmental conditions shape sensory systems and signaling traits, leading to divergent mate recognition signals that promote isolation upon secondary contact. Additionally, sexual selection drives rapid evolution of mating signals, as preferences for exaggerated traits in one population can quickly lead to assortative mating and reduced attraction to signals from divergent populations. Recent research from 2023 to 2025 on Drosophila species has revealed a polygenic basis for mate choice, where multiple genetic loci contribute to variation in courtship behaviors and discrimination, facilitating the buildup of isolation even in incipient stages. Prominent examples illustrate these principles across taxa. In collared and pied flycatchers (Ficedula albicollis and F. hypoleuca), dialect differences in song repertoires reduce hybridization rates, with males singing mixed song types showing up to 30% higher likelihood of interspecific mating compared to those with species-typical songs. Fireflies (Photinus spp.) exhibit species-specific flash patterns, where females respond only to the temporal and intensity cues of conspecific males, effectively preventing cross-attraction in sympatric assemblages. Similarly, in sympatric frog populations, such as those of Hyla species, advertisement calls diverge in frequency and pulse rate, enhancing acoustic discrimination and minimizing heterospecific responses. The strength of behavioral isolation is often amplified in sympatry through reinforcement, where selection against hybrid offspring intensifies mate discrimination to avoid maladaptive matings. This effect is quantified in laboratory discrimination assays, which measure female preference or male courtship vigor toward conspecific versus heterospecific stimuli, revealing isolation indices that can exceed 80% in reinforced populations compared to allopatric ones.

Mechanical Isolation

Mechanical isolation represents a prezygotic reproductive barrier where physical incompatibilities between mating structures or pollinator adaptations prevent successful sperm transfer or pollen deposition, thereby blocking between . This barrier arises from morphological divergences in genitalia, floral organs, or associated apparatus that render interspecific mating mechanically impossible or inefficient. In , it typically involves mismatched copulatory organs, while in , it stems from floral traits that exclude incompatible . The primary mechanism underlying mechanical isolation is the lock-and-key hypothesis, which proposes that reproductive structures evolve as complementary, species-specific morphologies to enforce , with genital evolution closely tracking overall species divergence. This process is accelerated by , promoting rapid co-evolution of male and female genitalia, often resulting in asymmetry and complexity that enhance specificity. Insect studies demonstrate this through elaborate genital designs that function as barriers, where even minor asymmetries disrupt coupling during attempted matings. A prominent example occurs in species, where genital differences impede copulation; for instance, in crosses between D. santomea females and D. yakuba males, a sclerotized spike on the male fails to fit female genital cavities, causing injuries and preventing sperm transfer. In the D. simulans , variations in posterior lobe reduce copulation duration and sperm displacement efficiency in interspecific matings compared to conspecifics. In , mechanical isolation manifests through pollinator specificity, such as mismatches between corolla length and bee tongue length; bee-pollinated flowers with specialized trigger mechanisms deposit only on bees possessing the precise tongue extension needed to access , excluding shorter-tongued species and preventing cross-pollination. Evidence for mechanical isolation derives from microscopic examinations of failed matings, revealing precise structural mismatches, such as size and shape incompatibilities in that halt intromission. These barriers play a key role in cryptic species complexes, like the Drosophila simulans , where subtle genital divergences maintain isolation despite morphological similarity, contributing to without overt external differences.

Gametic Isolation

Gametic isolation represents a prezygotic reproductive barrier where molecular and biochemical incompatibilities prevent the fusion of from different species, even after physical contact between and or and . This isolation arises from species-specific recognition mechanisms that block fertilization, ensuring that genetic exchange occurs only within compatible pairs and contributing to . In and , these barriers typically involve mismatches in surface proteins or glycoproteins that mediate gamete adhesion and activation, halting the process before formation. Key mechanisms of gametic isolation include surface protein mismatches that disrupt gamete recognition. In sea urchins, the sperm protein bindin binds to species-specific receptors on the egg vitelline envelope, such as EBR1, leading to rejection of heterospecific sperm due to even minor amino acid variations in bindin; as few as 10 changes can confer complete incompatibility. In mammals, the (ZP), an surrounding the egg, acts as a selective barrier through glycoproteins like ZP2 and ZP3, where a specific domain in ZP2 mediates binding to sperm proteins such as ZPBP, preventing cross-species adhesion and penetration. In plants, self-incompatibility loci, particularly S-RNase systems in families like , produce pistil-expressed ribonucleases that degrade in incompatible pollen tubes, extending to interspecific barriers where non-cognate S-haplotypes trigger similar rejection, blocking pollen tube growth toward the . These mechanisms highlight how of gamete recognition proteins enforces at the molecular level. Representative examples illustrate the efficacy of these barriers. In sea urchins such as Strongylocentrotus purpuratus and S. franciscanus, heterospecific sperm fail to agglutinate eggs due to bindin-receptor mismatches, resulting in near-complete isolation observed in broadcast spawning scenarios. In mammals, mouse (Mus musculus) sperm do not bind effectively to hamster (Mesocricetus auratus) ZP, attributed to divergent glycosylation patterns on ZP3 that alter sperm receptor specificity. For plants, interspecific pollinations in Arabidopsis thaliana and A. lyrata reveal pollen tube guidance failures, where species-specific defensin-like peptides (e.g., from the AT1G33415 cluster) attract only conspecific tubes, causing heterospecific pollen to arrest or misdirect in the pistil. Recent studies have also identified glycosylation differences in hybrid pollen, such as altered N-linked glycans on pollen wall proteins in Solanum hybrids, which impair tube elongation and contribute to isolation by disrupting pistil signaling. Detection of gametic isolation relies on experimental approaches that isolate molecular interactions. fertilization assays, such as mixing gametes from different and observing or rates under controlled conditions, quantify incompatibility; for instance, sperm-egg binding assays show zero fertilization success between closely related . Genetic mapping identifies recognition loci through crosses and QTL analysis, as in where genes map to chromosomes controlling attraction, or in where S-locus variants correlate with interspecific rejection rates. These methods confirm biochemical barriers without postzygotic effects.

Postzygotic Barriers

Hybrid Inviability

Hybrid inviability is a postzygotic reproductive barrier in which hybrid zygotes form successfully following fertilization but fail to develop properly, resulting in embryonic or larval death before reaching reproductive maturity. This failure arises from genetic incompatibilities between the diverging parental genomes, disrupting essential developmental processes such as , organ formation, or metabolic function. Unlike prezygotic barriers that prevent zygote formation, hybrid inviability manifests after , often leading to early lethality that reinforces species boundaries by curtailing . Several mechanisms underlie hybrid inviability, including mitochondrial-nuclear mismatches that impair cellular energy production. In these cases, the mitochondrial inherited maternally interacts incompatibly with nuclear-encoded proteins from the paternal , causing defects in respiratory chain complexes and , which halt embryonic development. For instance, in interspecific hybrids, homozygous mismatches in complex I subunits lead to complete embryonic arrest or juvenile mortality. Dosage imbalances in represent another critical mechanism, particularly affecting the heterogametic sex (e.g., XY males), where divergent regulatory elements disrupt X-chromosome dosage compensation, resulting in overexpression or silencing errors that trigger pathways. For example, in hybrids, such incompatibilities cause male inviability. In Xenopus frog hybrids, activation of the p53 pathway contributes to inviability through at the blastula stage. Prominent examples of hybrid inviability occur in and systems. In crosses between Rana catesbeiana females and Rana clamitans males, hybrid embryos typically progress to the exogastrula stage but fail to complete , leading to 100% mortality; however, inducing triploidy can rescue viability by restoring genomic balance. Similarly, in interspecies crosses, such as those between and , hybrid seeds frequently abort due to endosperm breakdown from parent-of-origin genomic imbalances, where the triploid receives unequal maternal and paternal contributions, triggering nutrient starvation and collapse. These cases highlight how inviability targets early developmental checkpoints to prevent hybrid establishment. Quantification of hybrid inviability often reveals low survival rates in controlled crosses, underscoring its role as a potent barrier. For example, in centrarchid interspecific , embryo-to-adult averages below 50% in many F1 crosses, with viability declining by approximately 3.13% per million years of parental . Recent genomic approaches have advanced of underlying loci; a 2025 study in used pooled sequencing of surviving F2 to map candidate inviability regions linked to chromosomal fusions, revealing polygenic contributions to lethality in over 20% of screened genomic intervals. These findings emphasize the cumulative driving inviability, with varying from near-zero in closely related taxa to partial rescue in manipulated experiments.

Hybrid Sterility

Hybrid sterility represents a key postzygotic barrier to , in which hybrid individuals survive to reproductive adulthood but fail to produce functional , rendering them infertile. This contrasts with hybrid inviability, where embryos or juveniles perish before maturity. The underlying causes typically involve genetic incompatibilities that impair or gamete maturation, ensuring that interspecific crosses do not propagate viable offspring. Several mechanisms contribute to hybrid sterility, often stemming from evolutionary divergence between parental genomes. One primary cause is resulting from mismatches, where differing numbers or structures between species prevent proper segregation during , leading to unbalanced s. For instance, failure of homologous chromosomes to pair (asynapsis) arrests meiotic progression, disrupting formation and recombination, which halts development. Additionally, hormonal disruptions in hybrids can alter endocrine signaling, affecting gonadal function and or production, as observed in cases where endogenous imbalances obstruct microspore development. Prominent examples illustrate these mechanisms across taxa. In mammals, mules—hybrids between (64 chromosomes) and donkeys (62 chromosomes)—possess an odd number of 63 chromosomes, causing incomplete pairing and meiotic arrest in spermatocytes, resulting in no viable sperm. In insects, hybrids between Drosophila simulans and D. mauritiana exhibit male sterility due to asynapsis and elements that distort segregation, leading to defective . Among plants, interspecific hybrids often display sterility from mismatches that induce aneuploid gametes and failed , reducing fertility in advanced generations. Hybrid sterility frequently shows asymmetry, disproportionately affecting the (e.g., males in mammals or males in ), a pattern briefly linked to without implying detailed evolutionary drivers. This sex bias arises from hemizygosity of the in hybrids, amplifying incompatibilities during in the heterogametic individuals.

Hybrid Breakdown

Hybrid breakdown is a postzygotic reproductive barrier characterized by the reduced viability, , or fitness in the progeny of fertile F1 hybrids, typically manifesting in the or backcrosses due to disruptive genetic interactions between parental genomes. Unlike F1-specific issues, F1 hybrids in such cases are generally viable and capable of producing offspring, but subsequent generations suffer from sterility, inviability, or as a result of recombination exposing incompatible gene combinations. This phenomenon arises from epistatic interactions, where alleles that function well within their native genetic backgrounds become deleterious when recombined. The mechanisms underlying hybrid breakdown primarily involve the breakdown of co-adapted complexes during in F1 hybrids, which shuffles alleles evolved in , leading to novel epistatic mismatches in F2 progeny. Recombination can unmask recessive deleterious alleles or create imbalances in dosage-sensitive interactions, such as those involving chromosomal rearrangements or Dobzhansky-Muller incompatibilities that only become apparent beyond the F1 stage. Additionally, ecological contributes, as recombinant hybrids often fail to thrive in either parental due to traits, such as suboptimal immune responses or . These processes highlight how hybrid breakdown enforces by penalizing of foreign alleles. A classic example occurs in (Oryza sativa) subspecies hybrids, where F1 plants are fertile, but F2 generations exhibit severe spikelet sterility linked to loci like hsa1, involving tightly linked genes that cause and abortion through allelic interactions. In house mice (Mus musculus), appears in immune function among F2 or backcross progeny from subspecies crosses, where incompatible alleles lead to dysregulated immune responses, increased , or heightened susceptibility to pathogens, reducing overall . Recent studies on monkeyflowers (Mimulus spp.), including work up to 2023, reveal F2 through and , driven by parent-of-origin effects and production failures that expose genetic conflicts in recombinant offspring. The implications of hybrid breakdown extend to limiting across boundaries, as the costs in later hybrid generations prevent the stable incorporation of beneficial alleles, thereby stabilizing distinct gene pools and accelerating . By acting as a "sink" for maladaptive recombinants, it reinforces reproductive isolation without relying on immediate F1 barriers, contributing to the long-term divergence of populations in or secondary contact.

Genetic and Molecular Mechanisms

Dobzhansky-Muller Incompatibilities

Dobzhansky-Muller incompatibilities (DMIs) arise when genetic changes accumulate independently in diverged populations, leading to negative epistatic interactions between s upon hybridization, resulting in postzygotic reproductive barriers such as hybrid inviability or sterility. In this model, an ancestral population with compatible alleles at interacting loci (e.g., genotype AABB) splits into isolated s; one lineage fixes a derived at the first locus (aaBB), which is or advantageous in its genetic background, while the other fixes a derived at the second locus (AAbb), similarly compatible within its own population. Upon secondary contact, hybrids carrying both derived alleles (aabb) exhibit dysfunction due to the incompatible interaction, while parental genotypes remain unaffected. This framework, first proposed by Bateson in 1909 and formalized by Dobzhansky in 1937 and Muller in 1942, explains how reproductive isolation evolves as a of without requiring selection against hybrids in pure . The model extends beyond two loci to multiple interacting genes, creating a "" where additional DMIs accumulate over time, progressively strengthening isolation between . Each new incompatibility adds to the epistatic network without imposing fitness costs on the originating populations, allowing divergence to proceed neutrally or adaptively within each . Formalized in and , this Bateson-Dobzhansky-Muller framework highlights how simple allelic substitutions can generate complex defects, with the number of loci involved scaling with . In polygenic scenarios, simulations and theoretical models demonstrate that DMIs build cumulatively, often involving dozens of loci to produce complete barriers. Empirical evidence for DMIs comes from quantitative trait locus (QTL) mapping in model organisms, which has identified specific hybrid incompatibility genes. In yeast (Saccharomyces species), QTL analyses of spores have pinpointed nuclear-mitochondrial interactions causing sterility, such as incompatibilities in genes, with multiple loci contributing to spore inviability. Similarly, in Drosophila species, QTL mapping has revealed epistatic pairs like Hmr and Lhr, where derived alleles interact to cause , and broader genomic scans confirm polygenic architectures underlying sterility. Recent genomic studies from 2020 to 2025, including high-resolution sequencing of hybrid zones, show that DMIs accumulate as polygenic traits, with QTL data indicating that hybrid defects often involve 10–50 interacting loci across the genome. DMIs provide a genetic explanation for both hybrid inviability and sterility, offering a for intrinsic postzygotic that incurs no adaptive cost in non- contexts, thus facilitating in allopatric populations. This model applies broadly to both phenomena, as seen in cases where allelic mismatches disrupt developmental pathways (inviability) or (sterility), and it underscores the role of in evolutionary . By resolving how incompatibilities emerge passively during or drift, DMIs highlight the genetic underpinnings of reproductive without invoking direct selection for barriers.

Chromosomal Rearrangements

Chromosomal rearrangements, such as inversions, translocations, and fusions, alter the physical structure of chromosomes and suppress recombination in heterozygous hybrids, often leading to the production of unbalanced gametes and reduced fertility.02187-5) These structural changes create barriers to by disrupting meiotic pairing and segregation, thereby contributing to reproductive isolation between diverging populations. The primary mechanism involves meiotic disruptions in hybrids, where heterozygosity for a rearrangement results in crossover suppression within the rearranged region, producing aneuploid gametes with duplications or deletions that cause spore inviability or reduced fertility. In yeast, for instance, inversion heterozygotes in Saccharomyces cerevisiae exhibit sterility due to high rates of aneuploid spores from unbalanced meiosis, effectively halting gene exchange between strains with differing karyotypes.00385-6) Such rearrangements also indirectly suppress gene flow by linking adaptive alleles or Dobzhansky-Muller incompatibility genes within non-recombining blocks, enhancing isolation over time. Prominent examples include paracentric inversions in Drosophila pseudoobscura, which contribute to hybrid male sterility by generating dicentric bridges and acentric fragments during meiosis, reducing sperm viability in crosses with related species like D. persimilis. In plants, chromosomal rearrangements distinguish hybrid sunflower species (Helianthus anomalus and H. deserticola) from their parents (H. annuus and H. petiolaris), where extensive inversions and translocations contribute substantially to F1 hybrid sterility, with nine of eleven QTL for pollen viability mapping to these rearranged regions and accounting for 87% of the phenotypic variance in pollen sterility. Recent studies on chromosome fusions, such as those in rockfishes (Sebastes spp.), demonstrate how multiple fusions repattern recombination rates and drive sympatric speciation by causing hybrid meiotic instability, with isolation strengthening as fusion number increases. Detection of these rearrangements typically relies on karyotyping to visualize gross structural differences and whole-genome sequencing to identify breakpoints and inversion spans at resolution. The strength of isolation conferred by rearrangements often correlates positively with the extent of chromosomal divergence and time since separation, as accumulated changes amplify meiotic errors in hybrids.

Microbial Contributions

Symbiotic microorganisms, particularly those inhabiting the gut or reproductive tracts of hosts, can contribute to hybrid incompatibility by eliciting mismatched immune responses or disrupting microbial transmission between parental lineages, thereby acting as extrinsic barriers to . These microbes influence postzygotic by altering through mechanisms such as dysbiosis-induced or failure in , which exacerbates incompatibilities beyond purely genetic factors. One key mechanism involves bacteria in , which induce cytoplasmic incompatibility by modifying sperm to block egg fertilization in uninfected or differently infected females, preventing successful formation at the gametic level. This process disrupts sperm-egg fusion through toxin-antidote systems encoded by bacterial genes, leading to embryonic lethality in hybrids. In , fungal symbionts such as endophytes can alter viability by shifting toward production at the expense of male development, potentially reducing cross-pollination success between divergent lineages. In Nasonia wasps, infections cause bidirectional cytoplasmic incompatibility, resulting in sterility or lethality in between species like Nasonia vitripennis and Nasonia giraulti, where mismatched infections prevent viable offspring. Similarly, in Drosophila species such as paulistorum, induces male sterility, which can be reversed by antibiotic treatment to eliminate the symbiont. Recent studies have demonstrated that transfers or germ-free rearing can rescue fitness in Nasonia by alleviating these microbial incompatibilities, highlighting the potential for experimental manipulation to overcome barriers. The evolutionary role of these microbes in accelerating reproductive isolation stems from their capacity for , which allows rapid spread across populations and fixation of incompatible strains, promoting without relying solely on host nuclear changes. Advances in symbiont have revealed how in microbes like enhances their manipulative capabilities, filling gaps in understanding extrinsic drivers of isolation.

Evolutionary Dynamics

Reinforcement and Selection

Reinforcement is the evolutionary process whereby natural or enhances prezygotic reproductive barriers between populations to reduce the production of low-fitness , typically occurring in areas of where hybridization is possible. This process evolves preferences, allowing individuals to discriminate against potential mates from divergent populations, thereby minimizing the energetic and costs associated with hybrid offspring. The selective pressure arises primarily from postzygotic barriers that render hybrids inviable, sterile, or ecologically maladapted, favoring traits that promote intraspecific mating. Mechanistically, can operate through direct selection against maladaptive , where individuals that avoid interspecific matings have higher , or indirectly via genetic correlations between loci and fitness traits. Theoretical models demonstrate that is most pronounced in hybrid zones, where and hybridization create "hotspots" of selection intensity, accelerating the of under conditions of low and strong hybrid disadvantage. For instance, one-allele mechanisms, involving a single genetic variant that reduces heterospecific matings across populations, facilitate more readily than two-allele scenarios requiring coordinated divergence. These models predict that strengthens barriers only when the benefits of outweigh the costs of choosiness, such as reduced mate availability. Empirical examples illustrate reinforcement in action. In fruit flies (Drosophila species), divergence in cuticular hydrocarbon pheromones has evolved under , enhancing female discrimination against heterospecific males in sympatric populations and reducing hybridization rates. Similarly, in cichlid fishes (Pundamilia spp.), female preferences for conspecific male nuptial colors—blue in deeper waters and red in shallower ones—have strengthened, limiting interspecific matings and production in sympatric zones. A 2025 meta-analysis of studies further links to reinforced , showing that plastic responses to divergent environments amplify prezygotic barriers during early stages. Evidence for reinforcement comes from comparative analyses across populations, which consistently reveal stronger prezygotic isolation, such as elevated mating discrimination, in sympatric versus allopatric pairs of species. This pattern holds in diverse taxa, including insects and vertebrates, supporting the prediction that sympatry generates hotspots of reproductive character displacement where barriers evolve rapidly in response to hybridization costs. Experimental studies in Drosophila confirm that such isolation can emerge within generations under controlled sympatry, provided hybrid fitness is sufficiently low.

Haldane's Rule

Haldane's rule, formulated by British evolutionary biologist in 1922, states that in the hybrid offspring of interspecies crosses, the —typically the male in mammals and or the ZW female in —is more likely to exhibit inviability or sterility compared to the homogametic sex. This pattern holds across diverse taxa with chromosomal sex determination, reflecting a key asymmetry in postzygotic reproductive isolation driven by sex-linked genetic factors. The rule was originally observed in animal hybrids and has since been documented in over 80 independent cases, underscoring its generality in processes. Several mechanisms explain the prevalence of Haldane's rule, primarily involving the unique properties of sex chromosomes in hybrids. The dominance theory posits that recessive deleterious alleles or Dobzhansky-Muller incompatibilities accumulate on the X (or Z) chromosome; in the heterogametic sex, hemizygosity unmasks these recessives, leading to dysfunction, whereas the homogametic sex (with two X chromosomes) benefits from masking by a compatible allele. Additionally, the faster rate of molecular evolution on the X chromosome—due to reduced recombination and exposure to selection—results in a higher density of hybrid-incompatible genes compared to autosomes. Failures in dosage compensation, where the heterogametic sex upregulates X-linked gene expression to match autosomal levels, can also disrupt hybrid development; mismatches between parental regulatory mechanisms exacerbate inviability or sterility in the affected sex. Classic examples illustrate the rule's manifestation across animal groups. In Drosophila species, such as D. melanogaster and D. simulans, hybrid males (XY heterogametic) are almost universally sterile due to X-linked incompatibilities disrupting , while hybrid females remain . In birds with ZW sex determination, the heterogametic females often show inviability or sterility in hybrids; for instance, crosses between species yield viable males but inviable or sterile females owing to Z-linked recessive incompatibilities. Mammalian hybrids, like those between horses ( caballus, 64 chromosomes) and donkeys ( asinus, 62 chromosomes), produce mules where males are invariably sterile from meiotic failure, while rare cases of female have been reported, aligning with XY heterogametic bias. Recent genomic studies from 2020 to 2025 have reinforced the rule's dependence on sex chromosomes by examining organisms lacking them. In and other organisms without differentiated sex chromosomes, such as many hermaphroditic flowering and yeast (Saccharomyces hybrids), hybrid dysfunction is typically symmetric, lacking the sex bias seen in heterogametic systems, as the absence of hemizygosity eliminates X-linkage dominance effects; however, these studies highlight analogous "sieve" patterns where hybrid adaptation is constrained by recessive incompatibilities, indirectly supporting the dominance mechanism's role in sex-linked cases.

Role in Speciation

Reproductive isolation serves as the critical endpoint in the speciation process, marking the point at which diverging populations become reproductively independent and evolve into distinct . This isolation arises through various evolutionary modes, including , where physical barriers like geographic separation prevent and allow independent evolution; , involving divergence along environmental gradients with limited dispersal; and , where populations diverge within the same geographic area due to ecological or behavioral factors. In each case, the accumulation of isolating mechanisms ensures that genetic exchange is sufficiently reduced to maintain lineage integrity, fulfilling the biological species concept proposed by . The buildup of reproductive barriers occurs through diverse mechanisms, such as in small populations, favoring local adaptations, and in , which can instantaneously create reproductive barriers by altering numbers. Recent studies from 2020 to 2025 highlight the role of in "porous genomes," where ongoing hybridization tests the strength of barriers, allowing only the most robust isolating mechanisms to persist and drive . For instance, in models of barrier evolution, can either erode weak barriers or reinforce strong ones, shaping the trajectory toward complete . , as a process accelerating sympatric by selecting against hybrids, exemplifies how selection can hasten this accumulation in overlapping ranges. Illustrative examples underscore this role. In Darwin's Galápagos finches (Geospiza spp.), has proceeded through the cumulative evolution of both pre- and postzygotic barriers, including beak morphology differences reducing mating success and hybrid inviability, leading to 18 recognized species despite occasional . Similarly, polyploid speciation in plants like Tragopogon mirus and T. miscellus, formed in the early via hybridization and doubling between introduced European species in , demonstrates instant reproductive isolation, as the allopolyploids are fertile among themselves but sterile with parental diploids. These cases show how isolation not only completes but also generates hotspots. Contemporary research addresses gaps in understanding by emphasizing incomplete isolation, where partial barriers allow hybrid speciation—new species arising from fertile hybrids—as seen in hybrid sunflowers (Helianthus spp.) adapting to extreme habitats. This perspective reveals that reproductive isolation predicts broader biodiversity patterns, such as higher speciation rates in lineages with rapid barrier evolution, challenging traditional views of strict isolation and highlighting its dynamic, multifaceted contribution to evolutionary diversification.

Interactions Among Mechanisms

Intrinsic vs. Extrinsic Barriers

Reproductive barriers in speciation are broadly classified into intrinsic and extrinsic types based on their dependence on environmental context. Intrinsic barriers are genetic incompatibilities that consistently reduce , such as sterility or inviability, irrespective of the ecological setting in which hybrids develop. These barriers arise from negative epistatic interactions between diverged genomes, as described by the Dobzhansky-Muller model, where alleles that function adaptively within their parental species become incompatible in hybrids. In contrast, extrinsic barriers are context-dependent, stemming from ecological mismatches where hybrids exhibit reduced due to to specific habitats, often resulting from divergent local selection pressures between parental populations. Mechanistically, intrinsic barriers primarily operate through Dobzhansky-Muller incompatibilities, which involve locus-specific genetic conflicts that disrupt essential cellular processes like or . Extrinsic barriers, however, emerge from divergent selection driving local or , leading to hybrids that perform poorly in the specialized environments of either parent. A 2025 meta-analysis of studies demonstrated that populations under divergent selection evolve stronger extrinsic reproductive isolation compared to those in uniform environments, highlighting the role of ecological divergence in accelerating these barriers. A classic example of an intrinsic barrier is observed in hybrid yeasts of the genus, where chromosomal rearrangements, such as inversions or translocations, cause meiotic sterility by preventing proper segregation, reducing viability to near zero regardless of growth conditions. For extrinsic barriers, lake and stream populations of threespine stickleback fish (Gasterosteus aculeatus) illustrate how hybrids suffer reduced survival and growth in parental habitats due to mismatched morphological traits—such as suited to open water versus streamlined forms for flowing streams—resulting in up to 50% lower fitness in non-native environments. Hybrids frequently encounter both intrinsic and extrinsic barriers simultaneously, compounding their fitness costs and reinforcing isolation, though the relative contributions vary by system. Recent hybrid zone studies from 2024–2025, including analyses in avian and plant systems, have quantified these effects using genomic and fitness assays, revealing that extrinsic barriers often play a key role early in divergence, while intrinsic incompatibilities accumulate later to stabilize boundaries. Gene flow can erode both barrier types by introducing adaptive alleles, but persistent ecological divergence tends to maintain extrinsic isolation more robustly than intrinsic genetic conflicts.

Effects of Gene Flow and Hybrid Zones

Hybrid zones represent narrow geographic regions where genetically diverged populations or species come into secondary contact, leading to interbreeding and the production of offspring, serving as natural laboratories for testing the strength of reproductive barriers under ongoing . These zones allow researchers to observe how partial isolation mechanisms function in real-time, revealing the interplay between and selection that either erodes or reinforces barriers to reproduction. In hybrid zones, —the transfer of genetic material across species boundaries—can erode weak reproductive barriers by allowing adaptive alleles to spread, particularly when dispersal rates exceed the strength of selection against hybrids. Tension zones, a common type maintained by a balance between dispersal and endogenous selection against unfit hybrids, often move over time in response to environmental gradients or demographic shifts, with permeable barriers facilitating asymmetric that favors one parental . Recent replicate studies across such zones, including those conducted in 2025, have identified specific barrier loci by sampling multiple parallel clines, demonstrating how genomic admixture highlights both parallel and divergent patterns in isolation architecture. A notable example is the European barn swallow (Hirundo rustica) hybrid zone, where behavioral barriers, such as sexual selection on plumage and song traits, limit the spread of introgressed genes, maintaining distinct lineages despite ongoing contact. In contrast, sunflower (Helianthus) hybrid swarms illustrate adaptive potential, as recombination in zones between species like H. annuus and H. petiolaris has generated novel genotypes that colonize extreme habitats, such as sand dunes, leading to the formation of reproductively isolated hybrid species. Advances in since 2020 have revealed that is commonplace even among recognized , with often occurring at low levels across permeable genomic regions while barrier loci resist exchange. These dynamics in hybrid zones predict outcomes like of prezygotic barriers to reduce maladaptive hybridization or, alternatively, when transgressive traits enable niche exploitation.

References

  1. [1]
    Speciation: The Origin of New Species | Learn Science at Scitable
    Reproductive Isolation: Genetically-based differences between populations which reduce or prevent genetic exchange between them (i.e., reproductive barriers).
  2. [2]
    Reproductive Isolating Mechanisms – Molecular Ecology & Evolution
    Definition: Mechanical isolation occurs when anatomical differences prevent successful mating between species. Example: Two species of insects might have ...
  3. [3]
    Ernst Mayr and the modern concept of species - PMC
    And finally, according to Ernst Mayr (9), “Species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated ...
  4. [4]
    Reproductive isolation - Understanding Evolution
    Speciation requires that the two incipient species be unable to produce viable offspring together or that they avoid mating with members of the other group.
  5. [5]
    Formation of New Species - OERTX
    Reproductive isolation can take place in a variety of ways. Scientists organize them into two groups: prezygotic barriers and postzygotic barriers.
  6. [6]
    What is reproductive isolation? - PubMed
    RI is a quantitative measure of the effect that genetic differences between populations have on gene flow.
  7. [7]
    The Process of Speciation – Introduction to Global Change
    In plants, new, reproductively isolated species may arise instantaneously, due to multiplication of the entire complement of chromosomes by a process known as ...<|control11|><|separator|>
  8. [8]
    What is reproductive isolation? - PMC - PubMed Central
    Reproductive isolation (RI) is a core concept in evolutionary biology and the basis by which biological species are defined.
  9. [9]
    Theodosius Dobzhansky on Hybrid Sterility and Speciation - PMC
    Dobzhansky's (1936) article in GENETICS represents the first concerted effort to work out the genetic changes producing a puzzling reproductive barrier: hybrid ...
  10. [10]
    Ernst Mayr and the modern concept of species - PNAS
    Apr 25, 2005 · Ernst Mayr played a central role in the establishment of the general concept of species as metapopulation lineages.
  11. [11]
    Genomics of plant speciation - ScienceDirect.com
    Sep 11, 2023 · Geographic isolation is thought to facilitate speciation by eliminating the possibility of gene flow between diverging lineages. If geographic ...
  12. [12]
    Reproductive isolation is a heuristic, not a measure: a commentary ...
    Sep 5, 2022 · Complete reproductive isolation, where gene flow is effectively zero, is regarded by some biologists as an important end point of speciation.
  13. [13]
    Reproductive Isolation - an overview | ScienceDirect Topics
    Reproductive isolation is defined as the combined effect of barriers to gene flow between divergent populations, encompassing both prezygotic isolation, ...Missing: seminal papers
  14. [14]
    How mechanisms of habitat preference evolve and promote ... - NIH
    Abstract. Habitat preference may promote adaptive divergence and speciation, yet the conditions under which this is likely are insufficiently explored.
  15. [15]
    Fruit odor discrimination and sympatric host race formation ... - PNAS
    pomonella is directly tied to host choice, the difference in host odor preference results in premating reproductive isolation between apple and hawthorn flies.Abstract · Sign Up For Pnas Alerts · Materials And Methods
  16. [16]
    Reproductive Isolation Among Deep-Water Cichlid Fishes of Lake ...
    However, cichlid species can naturally co-occur in narrow light spectrum habitats, such as turbid shallow lakes and the deep benthic zones of African rift lakes ...
  17. [17]
    Speciation dynamics and extent of parallel evolution along a lake ...
    Nov 3, 2021 · Here, we investigate differentiation trajectories and the extent of parallel evolution in East African cichlid fishes, which constitute ...
  18. [18]
    Divergent dynamics of sexual and habitat isolation at the transition ...
    Mar 13, 2024 · Speciation is often viewed as a continuum along which populations diverge until they become reproductively-isolated species.
  19. [19]
    Evolution of strong reproductive isolation in plants - Journals
    Jul 13, 2020 · Current research on speciation genomics strives to tackle two central questions in evolutionary biology: what is the origin and evolution of ...
  20. [20]
    Quantifying temporal isolation: a modelling approach assessing the ...
    Flowering time divergence can be a crucial component of reproductive isolation between sympatric populations, but few studies have quantified its actual ...
  21. [21]
    Temperature and photoperiod as environmental cues affect body ...
    Mar 1, 2017 · Seasonal changes in temperature and photoperiod are important environmental cues used by small birds to adjust their body mass (Mb) and ...
  22. [22]
    Meta-analysis reveals that phenotypic plasticity and divergent ...
    May 7, 2025 · A central tenet of speciation research is that reproductive isolation should increase with time as two populations or incipient species diverge.
  23. [23]
    Ecological and genetic divergence between two lineages of Middle ...
    Ecological prezygotic isolation could arise as a result of adaptation to contrasting habitats or temporal isolation [46]. ... Hyla arborea. Hereditas. 2000 ...
  24. [24]
    Temporal Separation and Speciation in Periodical Cicadas
    Feb 1, 2003 · Three distinct species with 17-year life cycles coexist in northern and Great Plains states (M. septendecim, M. cassini, and M. septendecula).
  25. [25]
    18.2B: Reproductive Isolation - Biology LibreTexts
    Nov 23, 2024 · Differences in breeding schedules, called temporal isolation, can act as a form of reproductive isolation. For example, two species of frogs ...
  26. [26]
    Ecological speciation by temporal isolation in a population of the ...
    Feb 10, 2017 · We conclude that this population is in the process of sympatric speciation, with temporal isolation being the most important direct barrier to gene flow.
  27. [27]
    Warmer springs increase potential for temporal reproductive ...
    Aug 14, 2023 · We found that warmer springs are associated with more temporal differentiation in flowering peaks among habitat patches, and less flowering ...
  28. [28]
    A perspective on sensory drive - PMC - PubMed Central
    Jul 6, 2018 · The basic premise of sensory drive is that variation in the environmental conditions under which signaling takes place can have profound effects ...
  29. [29]
    Sexual selection accelerates signal evolution during speciation in ...
    Sep 7, 2013 · We show that elevated levels of sexual selection are associated with more rapid phenotypic divergence between related lineages.
  30. [30]
    The genetic basis of incipient sexual isolation in Drosophila ...
    Jul 24, 2024 · We found that the genetic basis of incipient sexual isolation between DGRP males and Z females is highly polygenic and associated with the ...
  31. [31]
    Song similarity predicts hybridization in flycatchers - 2006
    May 12, 2006 · From the perspective of a male pied flycatcher, singing a mixed song type is associated with 30% likelihood of hybridization. This result, ...
  32. [32]
    Species-Specific Flash Patterns Track the Nocturnal Behavior of ...
    Jan 1, 2022 · Our study suggests that FI patterns may be a reliable species-specific luminous marker for monitoring the behavioral changes in a sympatric firefly population ...
  33. [33]
    Time and place affect the acoustic structure of frog advertisement calls
    Jul 26, 2024 · Often, these studies show that populations of sympatric frog species have calls that are more acoustically divergent than populations that ...
  34. [34]
    Reinforcement as an initiator of population divergence and speciation
    First, reinforcement can directly generate reproductive isolation between sympatric and allopatric populations. Specifically, reinforcement can generate the ...
  35. [35]
    [PDF] QUANTIFYING BEHAVIORAL ISOLATION BETWEEN CLOSELY ...
    Behavioral isolation is a common barrier that separates closely related ... infertile mating occurs ... reproductive isolation. Evolution 68:1511-1522 ...
  36. [36]
    The Molecular Mechanisms of Gametic Incompatibility in Invertebrates
    Gametic incompatibility is one of the mechanisms of reproductive isolation. It is based on species-specific molecular interactions that prevent heterospecific ...
  37. [37]
    A single domain of the ZP2 zona pellucida protein mediates gamete ...
    Jun 16, 2014 · The extracellular zona pellucida surrounds ovulated eggs and mediates gamete recognition that is essential for mammalian fertilization.
  38. [38]
    Compatibility and incompatibility in S-RNase-based systems - NIH
    Jul 28, 2011 · S-RNase-based self-incompatibility (SI) occurs in the Solanaceae, Rosaceae and Plantaginaceae. In all three families, compatibility is controlled by a ...
  39. [39]
    SEA URCHIN BINDIN DIVERGENCE PREDICTS GAMETE ...
    Using mitochondrial divergence as a proxy for time, we find that complete gamete incompatibility can evolve in approximately one and a half million years, ...
  40. [40]
    Gamete compatibility genes in mammals: candidates, applications ...
    Aug 30, 2017 · The second major layer is a thick glycoprotein layer called the zona pellucida (ZP) [7]. It often serves as a species-selective barrier for ...
  41. [41]
    A Species-Specific Cluster of Defensin-Like Genes Encodes ...
    Dec 18, 2012 · The flowering plant Arabidopsis thaliana has more than 300 defensin-like (DEFL) genes, which are likely to be involved in both natural immunity ...<|control11|><|separator|>
  42. [42]
    S-RNase Alleles Associated With Self-Compatibility in the Tomato ...
    Dec 5, 2021 · Our analysis focuses on 12 S-RNase alleles identified in SC species and populations across the tomato clade.Missing: gametic | Show results with:gametic
  43. [43]
    Bindin is essential for fertilization in the sea urchin - PNAS
    Aug 16, 2021 · Bindin isolated from the sperm acrosome results in insoluble particles that cause homospecific eggs to aggregate, whereas no aggregation ...
  44. [44]
    A lethal mitonuclear incompatibility in complex I of natural hybrids
    Jan 10, 2024 · Individuals homozygous for mismatched protein combinations do not complete embryonic development or die as juveniles, whereas those heterozygous ...
  45. [45]
    Aberrant X chromosome dosage compensation causes hybrid male ...
    This study showed that hybrid male inviability in Caenorhabditis arises from the dysregulation of X-chromosome dosage compensation, driven by the ...
  46. [46]
    Activation of P53 pathway contributes to Xenopus hybrid inviability
    May 15, 2023 · In fish and amphibians, many interspecific lethal hybrids arrest their embryonic development prior to gastrulation, which resembles the ...<|separator|>
  47. [47]
    Triploidy permits survival of an inviable amphibian hybrid - Elinson
    In the frog hybrid Rana catesbeiana female × Rana clamitans male, the embryo shows a characteristic development to an exogastrula which dies. This hybrid ...
  48. [48]
    Endosperm-based hybridization barriers explain the pattern of gene ...
    Jan 23, 2017 · This study shows that endosperm defects are sufficient to explain the direction of gene flow between the two wild species.Sign Up For Pnas Alerts · Results · Increased Ploidy Of A...
  49. [49]
    [PDF] TEMPO OF HYBRID INVIABILITY IN CENTRARCHID FISHES ...
    We found that hybrid embryo viability declined at mean rate of 3.13% per million years, slower than in most other taxa investigated to date. Despite measurement ...<|separator|>
  50. [50]
    Evolution of Hybrid Inviability Associated With Chromosome Fusions
    Feb 3, 2025 · We map hybrid inviability candidate loci by contrasting allele frequencies between F2 hybrids that survived until the adult stage with ...Missing: screens 2020-2025
  51. [51]
  52. [52]
  53. [53]
  54. [54]
    Genetic mechanisms of postzygotic reproductive isolation
    Dec 1, 2013 · Hybrid breakdown is defined as sterility or weakness observed in the F2 or later hybrid generations while the F1 hybrids grow normally with good ...
  55. [55]
    How Hybrid Breakdown Can Be Handled in Rice Crossbreeding?
    Oct 19, 2020 · Reduced hybrid viability and/or fertility segregating in F2 or later generations are referred to as hybrid breakdown (HB), in which recessive ...Introduction · The Genetic Basis of Hybrid... · Molecular Mechanisms... · Discussion
  56. [56]
    Two Tightly Linked Genes at the hsa1 Locus Cause Both F1 and F2 ...
    Feb 1, 2016 · Molecular mechanisms of hybrid breakdown associated with sterility (F2 sterility) are poorly understood as compared with those of F1 hybrid ...
  57. [57]
    A review on hybrid house mice - PMC - PubMed Central
    May 7, 2022 · The effect of hybridization in terms of immune defenses of hybrid mice ... hybrid breakdown by incompatibilities of such alleles. Altered ...
  58. [58]
    Understanding and overcoming hybrid lethality in seed and seedling ...
    In interspecies hybridization in Mimulus, F2 hybrid seedlings show lethality (hybrid breakdown) owing to a complete lack of chlorophyll production.
  59. [59]
    Evolutionary Genetics of Hybrid Incompatibility - Nature
    Dobzhansky (1937) and Muller (1942) independently formulated a model of how hybrid incompatibility could evolve. They both realized that hybrid inviability or ...
  60. [60]
    [PDF] GENETICS AND THE ORIGIN OF SPECIES
    GENETICS AND. THE ORIGIN OF SPECIES. BY. THEODOSIUS DOBZHANSKY. PROFESSOR OF ZOOLOGY IN. COLUMBIA UNIVERSITY. OXFORD BOOK COMPANY. CALCUTTA. NEW DELHI.
  61. [61]
    The Evolution of Hybrid Incompatibilities along a Phylogeny - NIH
    The Dobzhansky-Muller model of speciation posits that defects in hybrids between species are the result of negative epistatic interactions between alleles.
  62. [62]
    The evolution of postzygotic isolation: Accumulating Dobzhansky ...
    Aug 7, 2025 · According to Dobzhansky-Muller incompatibility theory, hybrids between species or populations with high genetic divergence are particularly ...<|control11|><|separator|>
  63. [63]
    Restoring fertility in yeast hybrids: Breeding and quantitative ... - NIH
    Sep 13, 2021 · QTL mapping in particular has shown to be a powerful tool for understanding the genetic basis of various complex traits and has been similarly ...
  64. [64]
    Two Dobzhansky-Muller genes interact to cause hybrid lethality in ...
    Nov 24, 2006 · The Dobzhansky-Muller model proposes that hybrid incompatibilities are caused by the interaction between genes that have functionally diverged in the ...Missing: evidence QTL yeast
  65. [65]
    The Impact of Chromosomal Rearrangements in Speciation
    Aug 21, 2023 · Various theoretical models have explored how chromosomal rearrangements (CRs) can be involved in speciation by reducing gene flow between ...
  66. [66]
    Chromosomal inversions and the reproductive isolation of species
    We suggest that inversions create linkage groups that cause sterility to persist between hybridizing taxa.
  67. [67]
    How chromosomal rearrangements shape adaptation and speciation
    This review examines the genetic causes and consequences of inversions as recombination suppressors and the role that recombination suppression plays in ...Missing: paper | Show results with:paper
  68. [68]
    Chromosome fusions repatterned recombination rate and facilitated ...
    Jan 30, 2023 · Early studies proposed that reproductive isolation is caused by abnormal chromosomal segregation in hybrid meiosis. However, experimental ...
  69. [69]
    Chromosomal Speciation in the Genomics Era - Frontiers
    Here, we revisit the early history of thinking about how chromosomal rearrangements (CRs) affect population and speciation processes.Missing: paper | Show results with:paper
  70. [70]
    The microbiome impacts host hybridization and speciation
    Oct 26, 2021 · Here, we survey and synthesize exemplar cases of how endosymbionts and microbial communities affect animal hybridization and vice versa.
  71. [71]
  72. [72]
    Fungal Symbionts as Manipulators of Plant Reproductive Biology
    Plants with a fungal symbiont produced more seeds and less pollen, shifting functional gender and switching from male-biased to female-biased sex allocation.Missing: viability isolation
  73. [73]
    Horizontal transmission maintains host specificity and ... - Nature
    Nov 16, 2023 · In host-symbiont systems, interspecific transmissions create opportunities for host switches, potentially leading to cophylogenetic ...Introduction · Discussion · Methods
  74. [74]
    Why Wolbachia-induced cytoplasmic incompatibility is so common
    Nov 7, 2022 · We argue that cytoplasmic incompatibility is pervasive because it enhances interspecific transmission and intraspecific persistence.
  75. [75]
    The What and Why of Research on Reinforcement - PMC
    Dec 14, 2004 · Reinforcement - a process that helps prevent interbreeding between hybridising populations - is an important and little understood mechanism ...
  76. [76]
    Speciation: The Strength of Natural Selection Driving Reinforcement
    Oct 6, 2014 · Speciation by reinforcement occurs when previously geographically separated populations come into secondary contact and hybridize. Individuals ...
  77. [77]
    [PDF] THE ROLE OF REINFORCEMENT IN SPECIATION: Theory and Data
    Sep 29, 2003 · Take, for example, a hypothetical case where a single allele causes females to prefer males that share their body size. If size had already ...<|control11|><|separator|>
  78. [78]
    the evolution of reproductive isolation beyond the first barriers
    Jul 13, 2020 · His research aims to determine the role of structural genomic rearrangements such as chromosomal fusion and fissions during the speciation ...Abstract · Is the evolution of strong... · The dynamics of reproductive... · Conclusion<|control11|><|separator|>
  79. [79]
    Female mating preference functions predict sexual selection against ...
    May 28, 2008 · Cichlid fish in African lakes have sustained high rates of speciation despite evidence for widespread hybridization, and sexual selection by ...
  80. [80]
    Reinforcement and other consequences of sympatry | Heredity
    Nov 1, 1999 · Researchers have repeatedly shown that sympatric species pairs tend to exhibit stronger species mating discrimination (sexual isolation) than ...
  81. [81]
    Degree of sympatry affects reinforcement in Drosophila - PubMed
    Specifically, premating isolation is strongest at intermediate degrees of sympatry. This result complements, rather than challenges, those of Yukilevich (2012).
  82. [82]
    Reinforcement can overcome gene flow during speciation in ... - NIH
    Dec 2, 2010 · Reinforcement is usually documented by observing a biogeographic pattern in which a reproductive isolating barrier is stronger in areas where ...
  83. [83]
    100 years of Haldane's rule - PMC - NIH
    Nov 10, 2022 · Haldane's rule is one of the 'two rules of speciation'. It states that if one sex is 'absent, rare or sterile' in a hybrid population, then that sex will be ...4. The Causes Of Haldane's... · 4.5. Dominance Theory · Figure 1
  84. [84]
  85. [85]
    Haldane's Rule: Genetic Bases and Their Empirical Support
    May 27, 2016 · Haldane's rule (HR), whereby the heterogametic sex suffers the most in terms of sterility, rarity, or absence following hybridization (Haldane 1922), is an ...The Genetic Mechanisms... · The Dominance Theory · Meiotic-Drive Theory
  86. [86]
    Beyond Haldane's rule: Sex-biased hybrid dysfunction for all modes ...
    Aug 19, 2024 · The genetic perturbation of a hybrid genome may disrupt the environmental response curve of sexual development to cause predictably skewed sex ...
  87. [87]
    Haldane's rule in the 21st century | Heredity - Nature
    Jan 12, 2011 · Haldane's Rule (HR), which states that 'when in the offspring of two different animal races one sex is absent, rare, or sterile, that sex is the heterozygous ( ...
  88. [88]
    Physiological aspects of sex differences and Haldane's rule ... - Nature
    Jul 1, 2022 · HR does not apply to the vast majority of angiosperm plants, as they are bisexual and have no sex chromosomes. Recently, it was revealed ...
  89. [89]
    Hybrid adaptation is hampered by Haldane's sieve - Nature
    Nov 28, 2024 · Together, these findings show that Haldane's sieve slows down adaptation in hybrids, revealing an intrinsic constraint of hybrid genomic ...Missing: Rule | Show results with:Rule
  90. [90]
    The importance of intrinsic postzygotic barriers throughout ... - Journals
    Jul 13, 2020 · Intrinsic postzygotic barriers can play an important and multifaceted role in speciation, but their contribution is often thought to be reserved to the final ...
  91. [91]
    Quantifying intrinsic and extrinsic isolating barriers across five ...
    Abstract. Understanding the relative roles of intrinsic and extrinsic reproductive barriers, and their interplay within the geographic context of diverging.
  92. [92]
    Comprehensive survey of condition-specific reproductive isolation ...
    May 26, 2015 · Chromosomal rearrangements as a major mechanism in the onset of reproductive isolation in Saccharomyces cerevisiae. Curr. Biol. 24, 1153 ...
  93. [93]
    Evolution of reproductive isolation in stickleback fish
    Extrinsic postmating barriers were common, and were stronger than intrinsic barriers in all systems, except the Lake–Stream system where the single estimated ...
  94. [94]
    Repeatable Selection on Large Ancestry Blocks in an Avian Hybrid ...
    Hybrid zones create natural tests of genetic incompatibilities by combining loci from 2 species in the same genetic background in the wild, making them useful ...Results · Introgression From Coastal... · Phenotype-Associated Loci...Missing: studies | Show results with:studies
  95. [95]
    Multiple hybrid zones involving four Cardamine species and their ...
    Sep 20, 2025 · Hybrid zones provide excellent opportunities to study evolutionary processes linked to interspecific gene flow, including introgression, ...Study Species · Plant Sampling · Hybrid Zones As A Window To...
  96. [96]
    A complex genomic architecture underlies reproductive isolation in a ...
    Feb 7, 2023 · Hybrid zones can be used as natural experiments to discover the processes that both facilitate and inhibit speciation. Mounting empirical and ...
  97. [97]
    Replicate geographic transects across a hybrid zone reveal ...
    May 7, 2025 · Hybrid zones provide a unique opportunity to use genetic admixture to identify traits and loci contributing to partial reproductive barriers ...
  98. [98]
    [PDF] Hybridization, Introgression, and the Nature of Species Boundaries
    Nov 16, 2021 · The notion that gene flow and reproductive isolation are characteristics of genome regions, not entire genomes, was already well established in.
  99. [99]
    Empirical study of hybrid zone movement | Heredity - Nature
    Jul 4, 2007 · Second, in a moving hybrid zone where barriers to gene flow are permeable, the effect of introgression will be most marked in the advancing ...
  100. [100]
    Sexual selection promotes reproductive isolation in barn swallows
    Dec 13, 2024 · Sexual signal traits form barriers to gene flow upon secondary contact. Barn swallows breed across nearly the entire North Hemisphere and ...
  101. [101]
    YEAR HISTORY OF A HYBRID POPULATION OF SUNFLOWERS ...
    Through introgression, species may acquire adaptive char- acters that allow them to colonize new habitats or increase their fitness in their existing niche more ...
  102. [102]
    Gene flow and introgression are pervasive forces shaping the ...
    Nov 10, 2022 · Here we analyzed the patterns of gene flow within and between species across >2600 bacterial species and >30,000 genomes. We identified which of ...
  103. [103]
    HYBRID SPECIATION IN WILD SUNFLOWERS1 - BioOne
    May 31, 2006 · We have previously shown that approximately 50% of the reproductive barrier between Helianthus species is caused by chromosomal rearrangements ...Hybrid Speciation: Theory · Ecological Selection · Conclusions And Future...