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Gene flow

Gene flow is the movement of genetic material between populations of the same species, occurring through the migration of individuals or the dispersal of gametes such as pollen or spores.^[1] This process, also referred to as genetic migration, transfers alleles—variant forms of genes—from one population to another, directly influencing the genetic composition of the recipient group.^[2] In evolutionary biology, gene flow serves as a key mechanism that homogenizes genetic variation across populations by counteracting the differentiating effects of natural selection and genetic drift.^[2] By introducing novel alleles into a population where they were previously absent, it enhances genetic diversity and can facilitate adaptation to new environments, such as through the spread of advantageous traits.^[1] Conversely, limited gene flow promotes genetic divergence between isolated populations, potentially leading to speciation when combined with local selective pressures.^[2] For instance, even a single migrant per generation can prevent substantial differentiation due to drift, maintaining connectivity in metapopulations.^[2] Mechanisms of gene flow vary by organism: in animals, it often involves physical migration of breeding individuals, while in plants, it relies on vectors like wind, water, or pollinators.^[1] Notable historical examples include interbreeding between Neanderthals and early modern humans, which transferred alleles influencing immune response, skin pigmentation, and metabolic traits like diabetes susceptibility into non-African human populations.^[1] In contemporary contexts, gene flow impacts conservation efforts, such as preventing inbreeding in fragmented habitats, and applied fields like agriculture, where it can either aid crop breeding or complicate management of genetically modified organisms.^[3] Overall, gene flow underscores the interconnectedness of populations, shaping evolutionary trajectories by balancing unity and diversity.^[2]

Fundamentals

Definition and Mechanisms

Gene flow refers to the transfer of genetic material, specifically alleles, between distinct populations of the same species, which can occur through the movement and interbreeding of individuals or the dispersal of gametes and reproductive structures. In population genetics, alleles are alternative forms of a gene that arise through mutations and contribute to genetic variation within a species, while populations are groups of interbreeding individuals that share a common gene pool—a collective set of all alleles present in the population. This transfer prevents populations from becoming genetically isolated, thereby reducing differentiation and promoting genetic cohesion across broader geographic ranges. The primary mechanisms of gene flow involve the physical dispersal of genetic material via migration of organisms or passive transport of reproductive elements. In animals, this often occurs through the migration and subsequent reproduction of individuals, such as birds traveling between distant habitats and mating with local populations, thereby introducing new alleles. In plants, gene flow is frequently mediated by pollen dispersal, which can be carried by wind, water, or pollinators; for instance, bees or birds transporting pollen between isolated plant populations facilitates cross-fertilization and allele exchange. Additionally, seed dispersal by wind, animals, or other vectors, like the wind-blown seeds of dandelions, enables the movement of entire zygotes to new areas where they can germinate and contribute to local gene pools. These processes require viable dispersal followed by successful reproduction and survival of offspring to establish gene flow. The concept of gene flow emerged in the field of population genetics during the early 20th century, building on foundational work by researchers like Ronald Fisher, J.B.S. Haldane, and Sewall Wright, who integrated it into models of evolutionary change. It was notably emphasized in Wright's 1931 paper on the shifting balance theory, where gene flow was described as a counterforce to genetic drift and selection in subdivided populations. In the context of the Hardy-Weinberg equilibrium, a cornerstone of population genetics established in 1908, gene flow represents a key violation of the model's assumptions—no migration or gene exchange—which would otherwise maintain constant allele frequencies in an idealized, closed population.

Importance in Evolution

Gene flow plays a pivotal role in maintaining genetic variation within populations by introducing new alleles through the movement of individuals or gametes, thereby counteracting the differentiating effects of genetic drift and reducing the risks associated with inbreeding.^[2] In small or isolated populations, genetic drift can lead to the random fixation or loss of alleles, diminishing diversity over time, while inbreeding increases homozygosity and the expression of deleterious recessive traits; gene flow mitigates these processes by replenishing genetic diversity, with even low levels of migration—such as one migrant per generation—sufficient to offset drift's impacts.^[2] This preservation of variation enhances a population's adaptability to changing environments, as diverse gene pools provide a broader reservoir of potential adaptive traits.^[4] Beyond maintaining variation, gene flow influences natural selection by facilitating the spread of beneficial alleles across populations, which can accelerate adaptive evolution and contribute to hybrid vigor, or heterosis, where offspring from diverse parents exhibit increased fitness.^[5] For instance, the influx of advantageous variants from neighboring populations can boost additive genetic variation, enabling faster responses to selective pressures like pathogens or environmental shifts, as observed in studies of parasite adaptation in natural systems.^[6] Hybrid vigor arises particularly when gene flow promotes outbreeding, leading to heightened heterozygosity and improved traits such as growth rate or disease resistance in the resulting progeny.^[7] However, the interaction between gene flow and selection is context-dependent, as moderate levels can enhance local adaptation without overwhelming endogenous evolutionary processes.^[8] Gene flow also bears directly on speciation by either homogenizing genetic differences among populations or, when restricted, permitting the divergence necessary for reproductive isolation to evolve. High rates of gene flow tend to unify allele frequencies across groups, preventing the accumulation of distinct genetic profiles that could lead to species formation.^[9] Conversely, low or absent gene flow allows populations to evolve independently under divergent selective pressures or drift, fostering genetic incompatibilities over time that reinforce barriers to interbreeding.^[2] This dynamic underscores gene flow's position as a key modulator in the speciation continuum, where its presence or absence shapes the trajectory of evolutionary divergence without invoking specific isolation mechanisms.^[9] Despite these benefits, gene flow introduces evolutionary trade-offs, as it can inadvertently spread maladaptive traits or dilute locally adapted genotypes, potentially hindering adaptation in heterogeneous environments. Incoming alleles mismatched to a recipient population's conditions may increase genetic load by favoring less fit variants, thereby counteracting the effects of local selection and imposing constraints on specialization.^[10] For example, migration from contrasting habitats can overwhelm beneficial local alleles, leading to reduced fitness in sink populations and exemplifying how gene flow balances connectivity against the risk of maladaptation.^[11] These trade-offs highlight gene flow's dual nature as both a promoter and limiter of evolutionary progress, depending on ecological context and migration intensity.^[12]

Quantification and Modeling

Measurement Techniques

Field-based techniques provide direct empirical estimates of gene flow by observing individual movements and dispersal events in natural populations. Mark-recapture studies, a cornerstone method, involve capturing, marking, and recapturing individuals to calculate migration rates between populations, offering insights into contemporary dispersal patterns.^[13] For instance, in fragmented habitats, these studies have quantified dispersal distances in species like butterflies and small mammals, revealing how landscape features influence migration success.^[14] In plants, pollen tracking techniques use fluorescent dyes or genetic paternity analysis to trace pollen dispersal, estimating gene flow via pollinator-mediated transfer.^[15] Such methods have demonstrated extensive pollen movement in wind-pollinated trees, where gene flow can span kilometers despite seed immobility.^[16] Molecular markers enable indirect inference of gene flow by analyzing genetic variation across populations, providing a scalable approach for large or elusive species. Microsatellites, highly polymorphic nuclear markers, are widely used to detect allele sharing that signals recent migration, often revealing higher gene flow than field observations suggest.^[17] Single nucleotide polymorphisms (SNPs), offering genome-wide resolution, allow precise estimation of differentiation levels, with thousands of loci improving accuracy in detecting subtle flows.^[18] Mitochondrial DNA (mtDNA), inherited maternally, tracks female-mediated gene flow but may underestimate total exchange due to its uniparental transmission.^[19] A key metric is the fixation index F_{ST}, which quantifies genetic differentiation as the proportion of total genetic variance attributable to differences among populations; low F_{ST} values indicate substantial gene flow homogenizing allele frequencies.^[13] Assignment tests leverage multilocus genotypes to assign individuals to source populations, thereby quantifying immigrant proportions and gene flow directionality. The software STRUCTURE employs Bayesian clustering to infer population structure and admixture, identifying the origins of individuals based on allele frequency profiles across markers like microsatellites or SNPs.^[20] This approach has been pivotal in studies of marine species, where it revealed ongoing gene flow despite apparent isolation. By estimating ancestry proportions, such tests distinguish migrants from residents, providing estimates of effective migration rates in systems with high genetic diversity.^[21] Despite their utility, these techniques face significant limitations that can bias gene flow estimates. Molecular methods, particularly those using F_{ST}, often reflect historical rather than recent gene flow, as genetic differentiation accumulates slowly and may not capture short-term changes in migration.^[13] Distinguishing contemporary from ancient exchange requires integrating multiple markers or time-series sampling, yet incomplete lineage sorting can confound interpretations.^[22] Sampling biases further complicate analyses; sparse or uneven sampling in low-density populations can inflate F_{ST} and underestimate flow, while overlooking rare long-distance events leads to underestimation of connectivity.^[23] Field-based approaches, though direct, suffer from logistical challenges in tracking elusive dispersers, potentially missing cryptic gene flow via undetected pathways.^[24]

Mathematical Models

Mathematical models of gene flow provide the theoretical foundation for understanding how migration influences genetic variation and differentiation among populations. One of the seminal frameworks is Sewall Wright's island model, which assumes a large number of equally sized subpopulations connected by symmetric migration at rate m, where each migrant contributes to the effective number of migrants per generation, Nm, with N being the effective population size per island.^[25] In this model, gene flow counteracts genetic drift, leading to an equilibrium level of population differentiation measured by F_{ST}, the fixation index representing the proportion of genetic variance attributable to differences between subpopulations.^[13] Approximating for small m, this simplifies to F_{ST} \approx 1 / (1 + 4N_e m), or rearranged, N_e m = (1 - F_{ST}) / (4 F_{ST}). This equation quantifies how gene flow (N_e m > 1) prevents substantial differentiation, while low gene flow allows drift to dominate.^[26]^[13] The continent-island model extends the island framework to asymmetric migration, typically from a large, genetically homogeneous continent (source) to a smaller island (recipient) at rate m, without back-migration. This setup is relevant for scenarios like colonization or peripheral populations. The change in allele frequency p on the island over one generation is given by the recursion p' = (1 - m) p + m p_c, where p_c is the allele frequency on the continent. Iterating this equation shows that the island frequency approaches p_c exponentially: p_t = p_c + (p_0 - p_c) (1 - m)^t, with the rate of convergence depending on m. At equilibrium under drift and mutation, differentiation measures like F_{ST} can be derived similarly, but adjusted for unidirectional flow, often yielding higher differentiation than symmetric models for equivalent m.^[25]^[27] For loci like microsatellites, which evolve under a stepwise mutation model (SMM), gene flow affects not just allele frequencies but the distribution of allele sizes, assuming mutations increment or decrement repeat units by one step with equal probability. In the SMM, proposed by Ohta and Kimura, the expected mean allele size in a subpopulation shifts under migration toward the source population's mean, analogous to the frequency recursion: the post-migration mean \mu' = (1 - m) \mu + m \mu_c, where \mu_c is the continental mean. The variance of allele sizes, which increases with time under mutation-drift balance, is reduced by migration homogenizing distributions across subpopulations, leading to an analog of F_{ST} called R_{ST} that incorporates size differences: R_{ST} \approx 1 / (1 + 8 N_e m) under an island model with symmetric stepwise mutations. This measure better captures differentiation at highly mutable loci, as migration mixes allele size spectra and slows divergence in both mean and variance.^[28] Applications of these models often involve coalescent theory simulations to evaluate complex scenarios beyond analytical tractability. Richard Hudson's ms software implements the coalescent process under the island model, generating genealogies for samples from multiple subpopulations with specified migration rates M = 4 N_e m, allowing inference of gene flow from simulated sequence data under neutral evolution. For intricate histories involving variable migration or barriers, approximate Bayesian computation (ABC) extends these simulations by comparing observed data summaries (e.g., F_{ST}) to those from coalescent-based priors, estimating posterior distributions of parameters like N_e m in non-equilibrium or hierarchical models.^[29]

Barriers and Isolation

Geographic Barriers

Geographic barriers, such as mountains, rivers, and oceans, physically separate populations and prevent interbreeding, leading to allopatric isolation where gene flow is restricted.^[30] These extrinsic obstacles reduce migration rates between subpopulations, allowing independent evolutionary trajectories to emerge over time.^[31] For instance, mountain ranges like the Andes have isolated Amazonian frog populations, resulting in distinct genetic clusters with minimal admixture.^[32] Vicariance events, where a once-continuous habitat is fragmented by geological processes, further exemplify how geographic barriers impede gene flow.^[33] Continental drift, as seen in the breakup of Gondwana, separated ancestral lineages across continents, promoting divergence without initial dispersal.^[34] Such events create long-term isolation, as evidenced by congruent phylogeographic patterns in co-distributed species like southern beeches and marsupials.^[35] The primary effect of these barriers is diminished gene flow, which fosters genetic divergence through processes like genetic drift and local adaptation.^[13] In Darwin's finches of the Galápagos Islands, oceanic distances between islands act as barriers, limiting migration and contributing to speciation among the 18 extant species, with genetic differentiation correlating to inter-island separation.^[36] This isolation has enabled adaptive radiations, where beak morphology evolved in response to varied food resources on separate islands.^[37] Quantifying the impact of geographic barriers often involves measuring genetic differentiation via F_ST values, which increase with physical distance due to isolation by distance.^[13] In equilibrium populations, pairwise F_ST estimates reflect reduced gene flow proportional to barrier strength, providing insights into migration rates without direct observation.^[38] For example, studies across fragmented landscapes show F_ST rising nonlinearly with separation, underscoring how barriers amplify divergence.^[39] Despite these barriers, gene flow can occasionally resume through rare long-distance dispersal events, such as rafting on floating vegetation or debris.^[40] In marine syngnathid fishes, rafting on kelp has facilitated transoceanic gene exchange, introducing alleles across otherwise isolated populations.^[41] Similarly, plant seeds and small invertebrates have colonized distant islands via rafting, occasionally countering vicariance-induced isolation.^[42]

Reproductive Barriers

Reproductive barriers represent intrinsic biological and behavioral mechanisms that impede gene flow between populations, even when their geographic ranges overlap, thereby facilitating divergence and speciation. These barriers are categorized into prezygotic, which prevent mating or fertilization, and postzygotic, which reduce the viability or fertility of hybrid offspring. By limiting interbreeding, such barriers maintain genetic distinctiveness and can drive evolutionary processes without reliance on physical separation.^[43] Prezygotic barriers act prior to zygote formation and include several key types that disrupt mate encounters or gamete fusion. Habitat isolation occurs when populations occupy different microhabitats within the same region, reducing encounters; for instance, one population may prefer rocky substrates while another favors sandy bottoms. Temporal isolation arises from differences in breeding timing, such as seasonal or diurnal variations in mating activity, preventing synchronous reproduction. Behavioral isolation involves divergent mating rituals or signals, like species-specific courtship dances or songs, that fail to elicit responses from potential mates of the other population. Mechanical isolation stems from morphological mismatches in reproductive structures, such as incompatible genitalia shapes, that hinder copulation. These mechanisms collectively minimize wasteful matings and are often the first line of defense against gene flow in sympatric conditions.^[44] Postzygotic barriers manifest after fertilization, typically through reduced hybrid fitness, which selects against interpopulation matings. Hybrid inviability results in embryos or offspring that fail to develop or survive to reproductive age due to genetic incompatibilities. Hybrid sterility produces viable but infertile adults, preventing further gene transmission; a prominent pattern is Haldane's rule, which observes that the heterogametic sex (e.g., XY males in mammals or ZW females in birds) is disproportionately affected by sterility or inviability in hybrids. This rule highlights the role of sex chromosomes in hybrid dysfunction, as hemizygosity exposes recessive incompatibilities. In response to such postzygotic costs, reinforcement can evolve, whereby natural selection strengthens prezygotic barriers—such as enhanced mate discrimination—to avoid producing low-fitness hybrids, thereby reducing gene flow in overlap zones.^[45]^[46] In sympatric contexts, where populations coexist without geographic separation, reproductive barriers are crucial for speciation by curtailing gene exchange amid potential interbreeding. African cichlid fishes in lakes like Victoria and Malawi exemplify this, where rapid radiations have produced hundreds of species through sensory-driven assortative mating and ecological specialization, leading to strong pre- and postzygotic isolation despite full range overlap. These barriers enable divergence via disruptive selection on traits like color patterns or feeding morphologies, fostering speciation without external isolation.^[47] The genetic underpinnings of these barriers, particularly postzygotic ones, often involve Dobzhansky-Muller incompatibilities, where alleles that are neutral or advantageous within their respective populations interact negatively in hybrids. Under this model, divergence in allopatry or local adaptation accumulates substitutions that become deleterious only in combination, such as mismatched proteins or regulatory elements, thus generating hybrid inviability or sterility without requiring adaptive conflict. This epistatic framework explains why even low levels of gene flow can be disrupted by a few key loci, reinforcing barriers over time.^[48]

Human Influences

Assisted Gene Flow

Assisted gene flow (AGF) refers to the intentional human-mediated movement of individuals, gametes, or seeds within a species' current range to enhance genetic diversity and facilitate adaptation to environmental changes, particularly climate change.^[49] This conservation strategy aims to mitigate inbreeding depression in isolated populations and introduce adaptive alleles to boost overall fitness, serving as a proactive tool for genetic rescue in endangered species.^[49] Unlike natural gene flow, which can be hindered by geographic barriers, AGF deliberately overcomes such isolation to maintain evolutionary potential in fragmented habitats.^[50] Key methods of AGF include translocation of live organisms, cryopreserved gametes for breeding, and seed transfer from seed banks to restoration sites.^[51] Translocation involves physically moving individuals from source populations to recipient sites, as seen in animal conservation, while seed banking supports plant programs by storing and deploying genetically diverse propagules for reintroduction.^[52] These approaches have demonstrated benefits such as increased heterozygosity and survival rates in small populations; for instance, genetic rescue efforts can reduce the frequency of deleterious alleles and substantially elevate fitness, with median increases of 45% in benign environments and 148% in stressful ones in the short term.^[53] A prominent example is the 1995 genetic rescue of the Florida panther (Puma concolor coryi), where eight individuals from Texas were translocated to the inbred south Florida population of fewer than 30 cats, resulting in hybrid offspring that comprised over 40% of the population by 2001 and contributed to a tripling of numbers to around 100 by 2010. As of 2025, the population has grown to around 200 individuals.^[54]^[55] Similarly, the 1995 reintroduction of 14 gray wolves (Canis lupus) from Canada into Yellowstone National Park introduced novel genetic lineages, enhancing gene flow across packs and maintaining high heterozygosity levels over subsequent generations.^[56] These interventions underscore AGF's role in reversing demographic declines through deliberate gene exchange. Despite its promise, AGF carries risks, including outbreeding depression, where maladaptive gene combinations from distant sources reduce hybrid fitness by disrupting locally adapted gene complexes.^[49] Ethical considerations involve balancing these risks against inbreeding threats, with guidelines emphasizing risk assessments, genetic monitoring, and prioritization of closely related sources to minimize negative outcomes.^[57] The International Union for Conservation of Nature (IUCN) provides frameworks for such translocations in its Guidelines for Reintroductions and Other Conservation Translocations, advocating assisted colonization variants for climate-vulnerable species while stressing ecological compatibility and long-term viability.^[58]

Genetic Pollution and Urbanization

Genetic pollution refers to the unintended introgression of genes from domesticated, genetically modified, or invasive organisms into wild populations, often leading to the erosion of native genetic integrity. This process is particularly concerning with genetically modified organisms (GMOs), where transgenes from crops such as rapeseed, rice, and sunflowers have escaped into wild relatives through hybridization, potentially altering ecological dynamics and reducing fitness in native species.^[59] For instance, genes from GMO crops can introgress into weedy relatives, conferring traits like herbicide resistance that may enhance invasiveness and disrupt local ecosystems.^[60] Urbanization exacerbates genetic isolation by fragmenting habitats, creating barriers that impede gene flow among populations. Roads, buildings, and impervious surfaces act as physical obstacles, significantly reducing dispersal in species like stream salamanders (Eurycea bislineata), where urban populations show higher genetic differentiation (F_ST = 0.110) compared to suburban or rural ones due to limited connectivity between streams.^[61] Edge effects from urban development further isolate remnants of natural habitat, increasing genetic drift and lowering effective population sizes (N_e ≈ 54 in urban vs. >1,000 in rural settings), which diminishes overall genetic variation despite short-term maintenance of diversity levels.^[61] The consequences of genetic pollution and urbanization-induced barriers include the loss of local adaptations and broader biodiversity erosion. In wild salmon populations, introgression from stocked hatchery strains—often non-local—has led to reduced reproductive success and phenotypic changes, swamping adaptive traits suited to specific environments and decreasing effective population sizes.^[62] Similarly, habitat fragmentation in urban landscapes heightens inbreeding risks and vulnerability to environmental stressors, potentially causing population declines and homogenization of genetic diversity across species.^[63] Mitigation strategies focus on restoring connectivity through urban planning initiatives, such as establishing green corridors to facilitate gene flow. Vegetated urban corridors and waterways have been shown to enable moderate dispersal in amphibians, countering isolation from roads and built environments in post-2020 studies.^[64] Conservation priority corridors, integrated into land-use policies, enhance genetic exchange by linking fragmented habitats, as demonstrated in frameworks aiming to protect 30% of land while allocating additional areas for wildlife movement.^[65] Recent analyses emphasize designing these corridors to bypass road barriers, promoting long-term population resilience in urbanizing regions.^[66]

Inter-Species Transfer

Hybridization

Hybridization involves the interbreeding of individuals from distinct species or subspecies, leading to the production of viable offspring that enable the exchange of genetic alleles between divergent genomes, thereby facilitating interspecies gene flow. This process integrates alleles from one lineage into another, potentially enhancing genetic diversity if the hybrids survive and reproduce.^[67] Hybrids can be classified as stable or unstable based on their reproductive viability. Stable hybrids maintain fertility and may establish independent lineages reproductively isolated from parental species, as exemplified by the sunflower species Helianthus anomalus, which arose from hybridization between H. annuus and H. petiolaris and occupies novel dune habitats. In contrast, unstable hybrids often suffer from sterility or low fitness due to genetic incompatibilities, restricting gene flow to limited introgression events. Evolutionary outcomes of hybridization include the introgression of adaptive traits that confer advantages in changing environments. For instance, hybridization between Neanderthals and modern humans introduced alleles enhancing immune responses in non-African populations.^[68] Hybridization also plays a pivotal role in adaptive radiation by generating transgressive variation—phenotypes beyond parental ranges—that allows colonizing populations to exploit vacant ecological niches and accelerate diversification.^[69] Barriers to hybridization often stem from partial reproductive isolation, which curtails gene flow despite occasional interbreeding. In annual sunflowers, chromosomal inversions and rearrangements between H. annuus and H. petiolaris create hybrid zones with reduced fertility, limiting allele exchange to specific genomic regions while maintaining species boundaries. These pre- and post-zygotic barriers, akin to those in intraspecific contexts, ensure that hybridization does not uniformly erode species distinctions. Detection of hybridization and resulting gene flow relies on genomic analyses that identify admixture signals in DNA sequences. Tools like ADMIXTURE software employ maximum likelihood methods to estimate ancestry proportions and detect introgressed segments from multilocus SNP data. Recent studies using such approaches have mapped Neanderthal introgression in modern human genomes, revealing ~2% archaic ancestry with selective retention of beneficial variants.^[69]

Horizontal Gene Transfer

Horizontal gene transfer (HGT), also known as lateral gene transfer, refers to the non-vertical movement of genetic material between organisms, distinct from parent-to-offspring inheritance, and serves as a significant mechanism of gene flow in microbial and some eukaryotic communities. Unlike vertical transmission, HGT allows for the rapid dissemination of adaptive traits across distantly related lineages, reshaping genomes and facilitating evolution in diverse environments. This process is particularly prevalent in prokaryotes, where it contributes to genetic diversity without requiring reproduction, but it also occurs in eukaryotes through specialized vectors.^[70] In bacteria, HGT primarily occurs via three mechanisms: conjugation, transformation, and transduction. Conjugation involves direct cell-to-cell contact through a pilus, enabling the transfer of plasmids or chromosomal DNA from a donor to a recipient bacterium. Transformation allows competent bacteria to uptake free DNA from the environment, incorporating it into their genome if it provides a selective advantage. Transduction is mediated by bacteriophages, which inadvertently package and deliver bacterial DNA during viral infection cycles. These mechanisms enable efficient gene exchange in dense populations, such as biofilms or soil matrices.^[71] In eukaryotes, HGT is less frequent but often involves transposon-mediated transfer, where mobile genetic elements excise from one genome and integrate into another via vectors like viruses or endosymbionts. Transposons facilitate the horizontal movement of genes across species boundaries, particularly in plants and invertebrates, by hijacking host machinery for dissemination. This process parallels aspects of hybridization in eukaryotes by introducing foreign DNA but occurs asexually without gamete fusion. Notable examples include the transfer of effector genes in fungi via giant transposons called "Starships," which drive rapid adaptation.^[72]^[73]^[74] The evolutionary significance of HGT is profound, as it accelerates adaptation by spreading beneficial alleles, such as antibiotic resistance genes among bacterial pathogens, which has led to global health challenges through mechanisms like plasmid conjugation. In endosymbiosis, HGT played a crucial role in the integration of ancient bacterial symbionts into eukaryotic cells; for instance, genes from the alphaproteobacterial ancestor of mitochondria were transferred to the host nucleus, enabling organelle function and eukaryotic complexity. This gene flow via HGT has been instrumental in major evolutionary transitions, from prokaryotic diversification to the origins of multicellular life.^[75]^[70] Representative examples illustrate HGT's role in natural ecosystems. In soil bacteria networks, fungal mycelia act as conduits for gene transfer by forming liquid films that enhance bacterial motility and conjugation, facilitating the exchange of carbon metabolic genes among diverse soil microbiota. In eukaryotes, bdelloid rotifers, which reproduce asexually, frequently acquire bacterial genes through HGT; for example, they have incorporated ice-binding protein genes from soil bacteria, conferring freezing tolerance in Antarctic environments. These cases highlight HGT's contribution to ecological resilience.^[76]^[77]^[78] Quantification of HGT events relies on metagenomic sequencing, which analyzes environmental DNA to identify anomalous gene phylogenies indicative of transfer, such as sequences with bacterial origins in eukaryotic genomes. Advanced co-barcoding methods in metagenomics enable precise detection of HGT within microbial communities by linking donor-recipient pairs. Mathematical models further estimate transfer rates, incorporating parameters like conjugation frequency and population density; for instance, stochastic models predict HGT rates in bacterial populations ranging from 10^{-6} to 10^{-3} per cell per generation, varying by environment and selective pressure. These tools underscore HGT's dynamic impact on gene flow.^[79]^[80]^[81]

Case Studies

Natural Examples

In wind-pollinated tree species such as oaks (Quercus spp.), pollen-mediated gene flow occurs over extensive distances, connecting populations across continental scales and preventing genetic isolation. Studies of European oak stands, including pedunculate oak (Quercus robur) and sessile oak (Quercus petraea), reveal that an average of 60% of pollen originates from outside local stands, with parentage analyses estimating up to 85% external pollen immigration. Pollen dispersal distances can exceed several kilometers, with mean values ranging from 16 meters in dense French forests to 5,400 meters in open Italian landscapes, following fat-tailed distributions that favor rare long-distance events. These dynamics ensure high contemporary genetic diversity and question the delineation of discrete population boundaries in such systems.^[82] Among animal systems, bird-mediated seed dispersal exemplifies gene flow in island ecosystems, where migratory and resident frugivores transport propagules between isolated habitats. In the Canary Archipelago, up to 1.2% of migratory birds arriving from Europe to sub-Saharan Africa carry viable seeds in their guts, enabling endozoochorous dispersal over oceanic barriers and introducing genetic material to remote islands like Alegranza. This process homogenizes genetic variation across archipelagos, as evidenced by shared alleles in dispersed plant populations, and supports connectivity in fragmented landscapes without human intervention. Similarly, in continental savannas, elephant (Loxodonta africana) migrations maintain expansive gene pools by facilitating male-biased dispersal and allele exchange among herds. Genetic analyses of savannah elephants in South Africa's Greater Kruger Biosphere indicate a single panmictic population with negligible structure, attributed to natural movements—especially post-fence removals mimicking historical connectivity—that introduce novel alleles and counteract drift, preserving diversity levels equivalent to effective population sizes of 500–700 individuals.^[83]^[84] Microbial gene flow in marine environments is prominently driven by ocean currents, which passively transport phytoplankton cells and genetic material across vast basins. In cosmopolitan species like the diatom Pseudo-nitzschia pungens, genomic data reveal significant genetic structuring with limited gene flow between distant populations, influenced by current-mediated dispersal that acts as a partial barrier despite its potential for connectivity, such as across salinity gradients. Basin-scale analyses of global plankton communities, including phytoplankton, demonstrate that currents impose a characteristic timescale of genetic mixing, with transport patterns explaining up to significant portions of biogeographic structure and enabling adaptive gene sharing in open-ocean settings.^[85]^[86] Fossil DNA provides evidence of ancient gene flow's long-term impacts, as seen in mammoth (Mammuthus spp.) lineages where hybridization shaped evolutionary trajectories. Ancient genomes from Siberian specimens dating back over 1 million years indicate gene flow from woolly mammoths (Mammuthus primigenius) into the North American Columbian mammoth (Mammuthus columbi), with admixture proportions of 57–62% woolly ancestry occurring around 420,000 years ago. This introgression, likely via migratory contacts during Pleistocene range expansions, introduced cold-adaptive alleles and influenced morphological divergence, highlighting how historical gene flow buffered populations against environmental shifts over millennia.^[87]

Human-Impacted Examples

Human activities have profoundly altered gene flow in various ecosystems, often with both positive conservation outcomes and unintended negative consequences. In fragmented forest habitats, wildlife corridors and reintroduction efforts aim to restore gene flow among isolated populations of European bison (Bison bonasus). For instance, programs in central-eastern Europe, including the establishment of subpopulations that function as connectivity corridors, seek to enhance genetic exchange between herds and reduce inbreeding risks, thereby improving overall population viability.^[88]^[89] These interventions demonstrate how human-managed connectivity can mimic natural dispersal patterns in anthropogenically divided landscapes. On the negative side, escaped aquaculture fish have facilitated invasive gene flow into wild stocks, threatening native genetic integrity. In Atlantic salmon (Salmo salar), escapees from fish farms interbreed with wild populations, leading to introgression that alters life-history traits and reduces fitness across the full life cycle of wild individuals.^[90] This gene flow, driven by frequent escapes—such as the 2025 Storm Amy event in Scotland where nearly 75,000 farmed salmon entered wild rivers—has contributed to declining wild populations by diluting locally adapted alleles.^[91] Such cases highlight the risks of aquaculture without robust containment measures. Climate change has prompted human interventions like assisted migration through coral reef transplants to bolster gene flow and resilience. A landmark 2021 study demonstrated successful assisted gene flow in the critically endangered elkhorn coral (Acropora palmata) by using cryopreserved sperm from heat-tolerant populations to fertilize eggs in vulnerable reefs, producing hybrid offspring with enhanced thermal tolerance and genetic diversity.^[92] This approach, applied in the Caribbean, has yielded the largest cryopreserved wildlife population to date, offering a scalable method to counteract rapid reef decline.^[93] Recent post-2020 research reveals how human transport amplifies gene flow in urban bee populations. Migratory beekeeping practices have increased genetic diversity and connectivity among honey bee (Apis mellifera) colonies by facilitating long-distance movement, potentially signaling broader homogenization trends in managed pollinators.^[94] Similarly, studies on the eastern carpenter bee (Xylocopa virginica) show elevated gene flow across urbanized landscapes compared to semi-natural areas, attributed to inadvertent human-mediated dispersal via vehicles and infrastructure.^[95] These findings underscore urbanization's role in reshaping pollinator genetics, with implications for ecosystem services like pollination.

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Pollen-mediated gene flow ensures connectivity among spatially ...
Dec 30, 2019 · Gene flow in plants via pollen and seeds is asymmetrical at different geographic scales. Orchid seeds are adapted to long-distance wind ...
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Wind-dispersed pollen mediates postglacial gene flow among refugia
The observed patterns led to the hypothesis that wind-pollinated plants are able to establish a highly efficient pollen-mediated gene flow among refugia during ...
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(PDF) Genetic estimates of population structure and gene flow
Aug 5, 2025 · Indirect methods using genetic markers are the primary measure of gene flow levels among interbreeding populations.Missing: software | Show results with:software
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an empirical comparison of microsatellite and SNP variation in ...
Jan 11, 2017 · Estimates of genetic differentiation (F ST) among populations derived from microsatellites were consistently higher than SNP-based estimates ...
[19]
[PDF] Guidelines for the analysis of population genetic data used in an ...
A low FST could mean either a relatively high rate of gene flow over time, or the recent cessation of gene flow altogether (because differentiation by genetic ...
[20]
STRUCTURE: applications, parameter settings, software
STRUCTURE both identifies populations from the data and assigns individuals to that population representing the best fit for the variation patterns found.
[21]
Inference of structure in subdivided populations at low levels of ...
Motivation: This article considers the problem of estimating population genetic subdivision from multilocus genotype data. A model is considered to make use of ...
[22]
Biases in Demographic Modeling Affect Our Understanding of ...
The systematic bias toward the choice of SC models when the real scenario generating the data had continuous gene flow, and the general overestimation of the ...
[23]
Sampling Issues - Evaluating Human Genetic Diversity - NCBI - NIH
For example, a sparse sample from a geographic area often cannot differentiate a single population with gene flow that is restricted by distance from fragmented ...
[24]
[PDF] Landscape approaches to historical and contemporary gene flow in ...
These approaches have been fostered by the development of new genetic markers and statistical methods, as well as an awareness that contemporary gene flow.
[25]
THE GENETICAL STRUCTURE OF POPULATIONS
BY SEWALL WRIGHT, University of Chicago. G a l h Lecture at University College, London, 1950. First let me acknowledge the great honour that I feel in being ...
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[PDF] migration / drift balance under wright's "island model"
Mar 15, 2001 · For example, some people have used Nm to measure "gene flow", for example between ... Fst=1over1+4Nm.html.
[27]
Lecture Summary 26 January 2001: Migration
Jan 26, 2001 · Continent-Island model. Examples: Mainland-island situations. The allele frequency p in the next generation will be. pi+1 = pi(1-m) + pMm. where ...
[28]
A measure of population subdivision based on microsatellite allele ...
It was found that, under the generalized stepwise mutation model, R( ST) provides relatively unbiased estimates of migration rates and times of population ...
[29]
Approximate Bayesian Computation Untangles Signatures of ...
Jan 27, 2022 · Using an Approximate Bayesian computation approach, we identify the model that best describes the history of gene flow between these taxa. This ...
[30]
Patterns of genomic divergence in sympatric and allopatric ...
... geographic isolation decreases gene flow and thus increases genetic distance between allopatric populations [4]. For example, mountains, rivers, oceans, and ...Read Mapping And Snp Calling · Genome Resequencing And... · Genomic Islands Of...
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Historical climate change and vicariance events contributed to the ...
May 20, 2024 · Vicariance events refer to the process of geographic isolation due to geo-climatic events, which leads to the habitat fragmentation of ancestral ...
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Biogeographic Analysis Reveals Ancient Continental Vicariance ...
Thus, continental drift provides a preliminary explanation, against which recent dispersal can be contrasted, using numerical methods. New algorithms ...
[33]
Vicariance Biogeography - Evolutionary Biology
Jul 23, 2025 · A foundational book proposing continental drift as an explanation to the distribution of life on Earth. ... Gene Flow · Genetics, Ecological ...
[34]
Darwin's Galápagos finches in modern biology - Journals
Apr 12, 2010 · Finally, the study adds to the growing body of evidence indicating that the speciation can occur despite gene flow between populations. The ...Missing: allopatric | Show results with:allopatric
[35]
Gene flow, ancient polymorphism, and ecological adaptation shape ...
It was suggested that these islands represent regions that are resistant to gene flow between incipient species and that drive the initial stages of ...
[36]
ISOLATION BY DISTANCE IN EQUILIBRIUM AND NON ... - PubMed
It is shown that for allele frequency data a useful measure of the extent of gene flow between a pair of populations is M∘=(1/FST-1)/4, ...
[37]
[PDF] GENETIC ISOLATION BY ENVIRONMENT OR DISTANCE
Gene flow patterns may align with geographic distance (IBD—isolation by distance), whereby immigration rates are inversely proportional to the distance between ...
[38]
Does rafting promote contemporary gene flow? Global and regional ...
Buoyant kelp such as Durvillaea antarctica have probably played a crucial role in favoring transoceanic, long distance dispersal by rafting, and genetic ...Missing: examples | Show results with:examples
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Asymmetrical gene flow in five co-distributed syngnathids explained ...
May 6, 2020 · Rafting dispersal has been hypothesized as the primary means of transport in these species [6,14,19], as they are among the most abundant fish ...
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Towards a More Nuanced Understanding of Long‐Distance Rafting ...
Feb 20, 2025 · In Nacella it is rare, probably extremely so, meaning gene flow is sufficiently low to allow allopatric speciation, but in Siphonaria rafting is ...
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Genetics and the Evolution of Prezygotic Isolation
Oct 16, 2023 · Prezygotic isolation includes barriers to gene flow between populations that occur before fertilization.
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Sep 24, 2022 · Prezygotic Barriers. There are several different types of prezygotic barriers, including temporal, habitat, behavioral, gametic, and mechanical.What is a Species? · Gene Pools · Reproductive Isolation · Prezygotic Barriers
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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 ( ...
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Coupling, Reinforcement, and Speciation | The American Naturalist
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[45]
Process and pattern in cichlid radiations – inferences for ...
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Dobzhansky-Muller model of hybrid incompatibility - Nature
A popular explanation is the Dobzhansky-Muller (DM) model of hybrid incompatibility. In the ancestral population, the genotype is AA BB.
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Assisted Gene Flow to Facilitate Local Adaptation to Climate Change
Nov 23, 2013 · However, AGF may cause outbreeding depression (especially if source and recipient populations have been long isolated) and may disrupt local ...<|separator|>
[48]
Evaluating Assisted Gene Flow in Marginal Populations of a High ...
Assisted Gene Flow, the movement of gametes from populations adapted to the changed environment to populations threatened by new conditions, has recently ...
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Assisted gene flow using cryopreserved sperm in critically ...
Sep 7, 2021 · Assisted gene flow (AGF) is a conservation genetic intervention to accelerate the adaptation of plant and animal populations to environmental ...Missing: definition | Show results with:definition
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[PDF] Restoration Seed Reserves for Assisted Gene Flow ... - Forest Service
This new type of seed orchard, a restoration seed reserve (RSR) targeting imperiled species, would incorporate into seed production the seed transfer concepts ...Missing: methods translocation
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Genetic rescue of Florida panthers reduced homozygosity but did ...
Aug 5, 2025 · In the 1990s, Florida panthers numbered <30 individuals suffering from inbreeding depression. In 1995, eight pumas from Texas were ...
[52]
[PDF] The genetic rescue of the Florida panther
Despite concerns, managers released eight Texas cats to locations across south Florida in 1995. Pairs of Texas females were released in close proximity to ...
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Heterozygosity of the Yellowstone wolves - PMC - NIH
They evaluated a variety of aspects of genetic diversity in the wolf population, which originated from 41 founders introduced in 1995 and 1996, and which has ...
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Assisted gene flow from outcrossing shows the potential for genetic ...
Dec 22, 2022 · Assisted gene flow may provide the variation needed to persist, and increase adaptive potential.
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[PDF] Guidelines for Reintroductions and Other Conservation Translocations
Assisted colonisation is most often viewed as a solution for species facing extreme threat from climate change, irrespective of their current conservation ...
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Scientists' warning on genetic pollution - PMC - NIH
Transgenes have 'escaped' from multiple genetically modified organisms, such as salmon, rapeseed, rice and sunflowers, into wild populations [27, 39–44].
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The application of GMOs in agriculture and in food production for a ...
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Urbanization reduces gene flow but not genetic diversity of stream ...
Urban fragmentation can create barriers within already complicated stream networks, limiting gene flow and creating more severe consequences than in simple ...
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Population-specific gene expression responses to hybridization ...
If this is also the case for Atlantic salmon, changes to gene expression following farm-wild introgression could result in the erosion of local adaptation.
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Severe consequences of habitat fragmentation on genetic diversity ...
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Limited evidence for genetic differentiation or adaptation in two ...
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Conservation priority corridors enhance the effectiveness of ... - Nature
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[63]
From science to impact: Conserving ecological connectivity in large ...
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[64]
Hybridization and Gene Flow | Learn Science at Scitable - Nature
Hybridization between species can allow alleles from one genetic background to integrate into another if favored by selection.
[65]
Neanderthal ancestry through time: Insights from genomes ... - Science
Dec 13, 2024 · Gene flow from Neanderthals has shaped genetic and phenotypic variation in modern humans. We generated a catalog of Neanderthal ancestry ...
[66]
Horizontal Gene Transfer and the History of Life - PMC - NIH
Conjugation is a one-way transmission mechanism of DNA from one cell to another via a “sexual pilus” by which DNA is transported. This mechanism has ...Horizontal Transfer And... · Evolution Of Gene Repertoire · Gene Tree/species Tree...
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Mechanisms of Horizontal Gene Transfer in Bacteria - PMC - NIH
Sep 6, 2018 · There are three “classical" methods of DNA transfer in nature: bacterial conjugation, natural transformation, and transduction (von Wintersdorff ...
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Horizontal transfer and evolution of transposable elements in ...
Mar 13, 2020 · Horizontal transfer of transposable elements (HTT) is an important process shaping eukaryote genomes, yet very few studies have quantified ...
[69]
Genetic exchange in eukaryotes through horizontal transfer
Jan 31, 2018 · Here, we will discuss the latest findings regarding HT among eukaryotes, mainly HT of transposons (HTT), establishing HTT once and for all as an ...
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Gene acquisition by giant transposons primes eukaryotes for rapid ...
Dec 6, 2024 · Giant transposons called “Starships” mediate horizontal gene transfer in fungi to drive repeated evolution and rapid adaptation.
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Horizontal Gene Transfer and Its Association with Antibiotic ... - NIH
Sep 18, 2019 · There are three main mechanisms of DNA transfer described in prokaryotic organisms: transformation, transduction, and conjugation (Figure 1) [25] ...1. Introduction · 2.2. Transduction · 2.3. Conjugation
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Mycelia as a focal point for horizontal gene transfer among soil ...
Nov 4, 2016 · Our study shows that the network structures of mycelia promote bacterial HGT by providing continuous liquid films in which bacterial migration and contacts are ...Missing: examples | Show results with:examples
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Dissecting the HGT network of carbon metabolic genes in soil-borne ...
Jul 7, 2023 · Here, we investigated the occurrence of horizontal gene transfer (HGT) and established the HGT network of carbon metabolic genes in 764 soil-borne microbiota ...
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Bacterial gene aids freezing tolerance of rotifers
Mar 4, 2025 · Bdelloid rotifers from an Antarctic algal community have acquired bacterial genes that encode a well-known family of ice-binding proteins called DUF3494 ...
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Detecting horizontal gene transfer with metagenomics co-barcoding ...
Feb 5, 2024 · In this study, to better identify horizontal gene transfer (HGT) in individual samples, we introduce a new co-barcoding sequencing system ...Results · Short-Reads Mngs Library... · Gene Prediction And...
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Current examining methods and mathematical models of horizontal ...
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Elevated rates of horizontal gene transfer in the industrialized ...
Apr 15, 2021 · We investigate the extent to which the rates and targets of horizontal gene transfer (HGT) vary across thousands of bacterial strains from 15 human populations.
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High Rates of Gene Flow by Pollen and Seed in Oak Populations ...
Gene flow predominantly occurs through dispersal of both seed and pollen, and the contemporary distribution of neutral genetic diversity across the landscape ...
[79]
Overseas seed dispersal by migratory birds - PMC - PubMed Central
Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in ...
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Conservation Genetic Assessment of Savannah Elephants ...
We investigated patterns of genetic variation in savannah elephants from the Greater Kruger Biosphere, with a focus on those in previously unstudied nature ...
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Limits to gene flow in a cosmopolitan marine planktonic diatom - PNAS
We show that gene flow between distant populations of the globally distributed, bloom-forming diatom species Pseudo-nitzschia pungens (clade I) is limited.<|separator|>
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Genomic evidence for global ocean plankton biogeography ... - eLife
Aug 3, 2022 · We found robust evidence for a basin-scale impact of transport by ocean currents on plankton biogeography, and on a characteristic timescale of ...
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Million-year-old DNA sheds light on the genomic history of mammoths
Aug 17, 2021 · ... evidence suggest that, compared to all other mammoths ... gene flow may have been unidirectional, from woolly mammoth into the Columbian mammoth.
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First Steps into the Wild – Exploration Behavior of European Bison ...
Fragmented habitats can be connected by establishing (sub)-populations that work as corridors, thus increasing population size and gene flow, minimizing the ...
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The Reintroduction Analysis of European Bison (Bison bonasus L ...
Jun 13, 2022 · The release and creation of new European bison nuclei in the wild creates the premises for natural contacts with the existing free populations ...1. Introduction · 2. Materials And Methods · 3. Results
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Introgression from farmed escapees affects the full life cycle of wild ...
Dec 22, 2021 · Escapees from fish farms pose a threat to wild stocks. ... Karlsson, Gene flow from domesticated escapes alters the life history of wild Atlantic ...
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Almost 75,000 farmed salmon in Scotland escaped into the wild after ...
Oct 14, 2025 · Scientists warn escaped farmed salmon could interbreed with wild fish, further endangering Scotland's fragile populations.
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Assisted gene flow using cryopreserved sperm in critically ... - PNAS
Sep 7, 2021 · When introducing alleles into a population, there is a risk of outbreeding depression (i.e., the introduction of alleles unsuitable for a local ...
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New Research Shows First Successful Demonstration of Assisted ...
Sep 10, 2021 · Large-scale assisted gene flow is a conservation intervention to accelerate species adaptation to climate change by importing genetic ...
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Honey Bee Diversity Is Swayed by Migratory Beekeeping and Trade ...
This may indicate recent increased gene flow and may signal an alarming trend toward greater movement of honey bees in the regions sampled. Long-term ...
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[PDF] Conservation insights from wild bee genetic studies - Rehan Lab
Feb 19, 2021 · The eastern carpenter bee, Xylocopa virginica, shows increased gene flow across human- altered environ- ments compared to semi- natural areas, ...Missing: post- | Show results with:post-