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Vernalization

Vernalization is the promotion of flowering in certain plants through exposure to prolonged periods of low, non-freezing temperatures, a physiological adaptation that synchronizes reproductive development with the onset of spring to avoid premature blooming during unpredictable autumn conditions.^[1]00528-3) This process, first experimentally demonstrated by Gustav Gassner in 1928 through studies on rye varieties requiring cold for floral induction, underpins the distinction between winter and spring cultivars in crops like wheat and barley, where winter types necessitate vernalization for timely heading.^[2] At the molecular level, vernalization often involves epigenetic silencing of floral repressors, such as the MADS-box transcription factor FLOWERING LOCUS C (FLC) in Arabidopsis thaliana, achieved via cold-induced histone modifications that maintain repression even after return to warmer conditions, exemplifying a stable "memory" of winter.^[3]^[4] While the phenomenon enhances agricultural predictability and yield stability in temperate regions, historical applications were marred by pseudoscientific extensions under Trofim Lysenko in the Soviet Union, who erroneously claimed heritable transmission of vernalized states, leading to famines through misguided policies that suppressed genetic evidence for fixed varietal differences.^[5] Contemporary research prioritizes quantitative trait loci like VRN1 in cereals, enabling breeding for climate-resilient varieties amid shifting winters.^[6]

Definition and Principles

Core Process and Requirements

Vernalization is the physiological process whereby plants exposed to prolonged cold temperatures acquire competence to initiate flowering upon return to warmer conditions, thereby aligning reproduction with favorable spring environments.^[7] This requirement evolved in temperate species to prevent autumn flowering that would expose developing seeds to winter frost.^[8] The core environmental cue involves exposure to temperatures typically ranging from 0°C to 10°C, with species-specific optima; for example, Arabidopsis thaliana achieves vernalization at 0–6°C, while winter wheat (Triticum aestivum) responds to temperatures below 9°C.^[9]^[10] Durations of 4 to 12 weeks are generally required to saturate the response, varying by genotype and conditions—such as 40 days at 11°C or longer at cooler temperatures in wheat cultivars.^[11]^[12] This process primarily operates in winter annuals, biennials, and rosette-stage perennials including wheat, Arabidopsis thaliana, and Brassica species, distinguishing it from photoperiodic responses to day length or autonomous maturation pathways.^[13] Vernalization treatments are effective when applied to imbibed seeds, seedlings, or vegetative rosettes, but efficacy diminishes in later reproductive stages.^[8] The induced flowering competence demonstrates mitotic stability, maintained across cell divisions without ongoing cold, as evidenced by experiments where vernalized seedlings or apices, upon warming, exhibit accelerated flowering under permissive photoperiods or autonomously.^[14]^[8] This stability ensures the plant's progeny revert to requiring cold for vernalization, preventing heritable epigenetic shifts.^[8]

Distinction from Other Flowering Cues

Vernalization promotes flowering competence through prolonged exposure to cold temperatures, typically 4–10°C for 4–12 weeks depending on species, inducing a heritable epigenetic state that persists post-cold, in contrast to photoperiodism, which senses day length via phytochrome and cryptochrome photoreceptors to activate floral integrators like FLOWERING LOCUS T (FT).^[15]^[16] This cold-specific memory enables plants to anticipate spring conditions without reliance on light duration cues, as demonstrated in Arabidopsis accessions where vernalized plants flower under non-inductive short days, unlike non-vernalized controls.^[17] Gibberellin signaling accelerates flowering in long-day or rosette species by upregulating SOC1 and LFY independently of cold, but vernalization operates through distinct chromatin-based repression of inhibitors, showing causal separation; for instance, gibberellin-deficient mutants still vernalize effectively, while vernalization-deficient lines respond to exogenous gibberellins only partially.^[18]^[19] Age-related flowering, via the autonomous pathway that reduces repressor transcripts like FLC over developmental time without environmental input, contrasts with vernalization's acute response to winter cold, ensuring the latter's specificity to seasonal risk assessment rather than mere maturation.^[17]^[20] In temperate cereals such as wheat, cold induction of VRN1 epigenetically silences repressors like VRN2, overriding inhibition of FT homologs and permitting flowering competence even under short-day conditions that would otherwise delay it in winter varieties, highlighting vernalization's partial independence from photoperiod integration.^[21]^[22] This distinction underscores vernalization's role in aligning reproduction with reduced frost probability after prolonged cold, a adaptation evident in fossil and genetic evidence from early temperate grasses where cold sensing predates photoperiod fine-tuning.^[23]^[24]

Historical Context

Pre-20th Century Observations

Farmers in ancient agricultural societies, particularly in the Fertile Crescent where Pooideae grasses like emmer wheat and barley were domesticated around 10,000 years before present, empirically selected for crop varieties capable of overwintering through cold exposure, which promoted robust spring flowering and higher yields compared to those sown later without such conditioning.^[23] This distinction between autumn-sown "winter" cereals, which relied on prolonged cold to transition to reproductive growth, and spring-sown varieties that flowered promptly without it, formed the basis of sowing practices in temperate regions, enhancing survival and productivity under seasonal climates.^[5] By the early 19th century, European observers formalized these observations, debating whether spring and winter wheat represented fixed varieties or environmentally induced forms. In his 1800 publication A New Farmer's Calendar, John Lawrence noted that winter wheat, when sown in autumn, benefited from cold to prevent premature heading and ensure timely maturation, while spring wheat lacked this necessity.^[5] Such knowledge guided crop management across Europe, where winter varieties yielded up to 20-30% more under cold-conditioned regimes than unexposed plantings.^[5] Experimental quantification emerged in the 1830s, predating formal vernalization theory. In 1837-1838, Colonel Abbott tested Flint winter wheat by soaking seeds, exposing them to chilling conditions, and sowing in spring; the treated plants flowered earlier and resembled spring varieties, demonstrating cold's role in hastening reproductive development without altering inherent varietal traits.^[5] These trials, reported in agricultural periodicals like The Monthly Genesee Farmer, confirmed farmers' empirical insights that controlled cold exposure could accelerate flowering in winter cereals by 4-6 weeks, influencing yield predictions and sowing strategies.^[5]

20th Century Formulation and Key Experiments

Gustav Gassner conducted foundational experiments in 1918, demonstrating that winter cereals such as rye and wheat require exposure to low temperatures shortly after germination to promote timely flowering. By subjecting imbibed seeds to near-freezing conditions (approximately 0°C) for extended periods, Gassner observed that treated plants flowered earlier and more uniformly compared to untreated controls, establishing cold as a critical physiological trigger for floral competence in these species.^[2]^[25] Building on Gassner's findings, Trofim Lysenko coined the term "jarovization" (later translated by him as "vernalization" from the Latin vernalis, meaning "of spring") in his 1928 publication detailing trials on winter wheat seeds in the Soviet Union. Lysenko's method involved moistening dry seeds to initiate imbibition (typically 50 parts water per 100 parts dry matter) followed by chilling at low temperatures for several weeks, which induced heritable flowering competence in the treated generation, enabling spring planting to yield crops as if naturally vernalized over winter. These experiments emphasized the necessity of moisture during cold exposure to activate the response, distinguishing it from mere dry storage.^[25]^[5] In the 1930s, international researchers replicated and validated vernalization effects in cereals, confirming that prolonged cold (often 4–6 weeks at 0–5°C) on imbibed seeds or young seedlings accelerated ear emergence and flowering in winter varieties of wheat and barley, independent of photoperiod. For instance, experiments in the United Kingdom demonstrated marked advancement in heading dates for treated winter barley and wheat, aligning with Russian results and extending applicability to field conditions.^[26]^[25] The vernalization response is physiological rather than an inheritance of acquired traits, as the induced flowering competence is not transmitted to subsequent generations; progeny of vernalized plants revert to requiring their own cold exposure if they are winter types. This non-transmissibility, observed consistently in cereal experiments, refutes claims of permanent genetic change and underscores the effect's reset between generations.^[27]^[28]

Influence of Political Ideologies on Research

In the Soviet Union from the early 1930s onward, Trofim Lysenko elevated vernalization—initially a technique to hasten seed germination through cold exposure—as a cornerstone of his agrobiological program, asserting that it induced heritable physiological changes in plants akin to Lamarckian inheritance of acquired traits.^[29] Lysenko rejected Mendelian genetics as incompatible with dialectical materialism, framing vernalization as evidence that environmental modifications could permanently alter plant heredity without genetic mutation, a claim empirically unsupported and contradicted by controlled breeding experiments showing no intergenerational transmission of such traits.^[30] This ideological alignment with Soviet Marxism-Leninism secured state patronage, including Stalin's personal endorsement in 1935, enabling Lysenko to dominate agricultural academies and enforce vernalization in collective farms despite inconsistent yields from rushed implementations.^[31] Political suppression under Lysenkoism dismantled genetic critiques of vernalization's limits, purging rivals like Nikolai Vavilov in 1940 and banning Mendelian research, which prioritized unverified Lamarckian hypotheses over data from heritability tests.^[32] Soviet policies mandating widespread vernalization from 1931 contributed to agricultural inefficiencies, with overoptimistic promises of doubled harvests failing amid poor adaptation to local conditions and exacerbating post-collectivization grain shortfalls through the 1940s.^[33] Official Soviet reports inflated successes while concealing failures, reflecting institutional bias toward ideology over empirical falsification, as dissenting data risked accusations of sabotage.^[34] Western researchers, unencumbered by such orthodoxy, affirmed vernalization's utility for synchronizing crop cycles—evidenced by British trials in the 1930s yielding 10-20% wheat yield increases under controlled conditions—but decoupled it from heritability claims, fostering incremental advances in plant physiology via genetic crosses that highlighted stable varietal differences.^[35] This apolitical empiricism contrasted sharply with Soviet prioritization of political utility, where vernalization symbolized class struggle against "formal genetics," delaying integrative genetic models until Lysenko's ouster in 1964 allowed partial recovery, though Soviet contributions to vernalization's molecular underpinnings remained minimal compared to post-1950s Western efforts.^[36]

Molecular Mechanisms

Epigenetic Repression of Flowering Repressors

Prolonged cold exposure during vernalization triggers the recruitment of Polycomb Repressive Complex 2 (PRC2) to chromatin regions of flowering repressor loci, initiating a cascade of histone modifications that enforce stable transcriptional silencing. PRC2, composed of core subunits such as CURLY LEAF (CLF), SWINGER (SWN), and FERTILIZATION INDEPENDENT ENDOSPERM 2 (FIE), deposits trimethylation on histone H3 at lysine 27 (H3K27me3), a hallmark repressive mark that compacts chromatin and inhibits RNA polymerase access. This H3K27me3 modification spreads bidirectionally from nucleation sites, often linked to antisense transcripts like COOLAIR, ensuring domain-wide repression without requiring DNA sequence alterations.^[37]^[38]^[39] The repression exhibits dose-dependency, where the duration and intensity of cold directly correlate with the extent of H3K27me3 accumulation and the degree of gene silencing, mimicking a quantitative "counter" of winter severity to calibrate flowering timing. Shorter cold periods yield partial repression with incomplete H3K27me3 spreading, while extended exposure (typically 4-6 weeks at 4-5°C) achieves near-complete silencing in a stochastic, allele-specific manner across cell populations. This progressive buildup allows plants to integrate cumulative cold signals, accelerating flowering proportionally upon return to warm conditions through reduced repressor output. Mathematical models describe this as a probabilistic switching of loci from active to silenced states, with repression strength scaling logarithmically with cold duration to buffer against environmental variability.^[40]^[41] Epigenetic stability arises from the self-reinforcing propagation of H3K27me3 during subsequent mitotic divisions post-vernalization, independent of ongoing cold signals or de novo transcription. During the non-dividing cold phase, initial nucleation establishes the mark via cold-responsive factors, creating a heritable template that PRC2 reads and amplifies in progeny cells under warmer temperatures. Verification through chromatin immunoprecipitation confirms H3K27me3 persistence correlates with flowering competence, while bisulfite sequencing reveals auxiliary DNA hypermethylation at some loci reinforces but does not initiate the core histone-based memory. This mechanism ensures mitotic inheritance without reliance on cellular proliferation during cold, providing causal fidelity to seasonal cues.^[42]^[43]^[44]

Key Genes and Pathways in Arabidopsis

In Arabidopsis thaliana, vernalization primarily acts through epigenetic repression of the floral repressor FLOWERING LOCUS C (FLC), enabling competence to flower upon return to warm conditions. Prolonged cold exposure induces VERNALIZATION INSENSITIVE 3 (VIN3), an RNA helicase that facilitates chromatin compaction at the FLC locus by promoting histone deacetylation and H3K27 trimethylation via Polycomb Repressive Complex 2 (PRC2).^[45] PHD-finger proteins such as VIL1 (VIN3-INTERACTING PROTEIN 1), part of the VIN3-PRC2 complex, bind H3K4me3-marked regions and recruit PRC2 to initiate stable silencing during cold.^[46] This repression is quantitative, depending on cold duration, typically requiring 4-6 weeks at 4°C for full effect in winter annual accessions.^[47] Post-vernalization, VRN1, encoding a MADS-domain transcription factor homologous to APETALA1, maintains FLC silencing through warm temperatures by counteracting reactivation mechanisms, as evidenced by vrn1 mutants showing incomplete FLC repression and delayed flowering after cold treatment.^[48] VRN1 interacts with PRC2 components like VRN2 (a Su(z)12 homolog) to sustain H3K27me3 marks, ensuring epigenetic memory unique to Brassicaceae species.^[47] This FLC-VRN1 axis distinguishes Arabidopsis vernalization from cereal pathways, where orthologous genes regulate different repressors.^[49] The autonomous pathway integrates with vernalization by constitutively repressing FLC via genes such as FCA (RNA recognition motif protein) and FVE (histone-binding protein), which promote alternative polyadenylation and histone deacetylation at FLC.^[50] Mutants like fca and fve exhibit elevated FLC expression and late flowering under long days, but vernalization bypasses this delay by directly silencing FLC, confirming the pathway's causal role in modulating cold responsiveness.^[51] Genetic screens in the 1990s identified these mutants, demonstrating that autonomous regulation sets a baseline FLC level, which vernalization further reduces for flowering promotion.^[52] Age-dependent competence to vernalize involves the miR156/miR172 module, where high juvenile miR156 levels repress SQUAMOSA PROMOTER-BINDING PROTEIN-LIKE (SPL) transcription factors, delaying miR172 accumulation and thus reducing sensitivity to cold-induced FLC repression.^[53] As plants age, declining miR156 allows SPLs to promote miR172, which targets APETALA2-like repressors, enhancing vernalization efficacy; this mechanism, conserved from studies in related Brassicaceae like Cardamine flexuosa (2013), modulates cold sensitivity in Arabidopsis juveniles versus adults.^[54]^[55]

Variations in Cereals and Other Species

In temperate cereals such as wheat (Triticum aestivum) and barley (Hordeum vulgare), vernalization induces flowering through the upregulation of VRN1, a MADS-box transcription factor that establishes floral meristem identity and overrides the winter growth habit.^[22] This process coincides with the repression of VRN2, a CONSTANS-like zinc finger gene that acts as a long-day flowering inhibitor and maintains vegetative growth prior to cold exposure.^[56] The VRN3 locus, orthologous to FLOWERING LOCUS T (FT), serves as an integrator of vernalization and photoperiod cues, transmitting the cold-promoted signal from leaves to the shoot apex to activate VRN1 under inductive long-day conditions.^[57] A fourth locus, VRN-D4 on the short arm of chromosome 5D, contributes to vernalization diversity via a ~290-kb insertion from chromosome 5AL, which originated in ancient South Asian wheat populations and facilitated the transition to spring sowing by reducing cold requirements.^[6] Regulatory mutations at VRN-D4 exhibit epistatic interactions with VRN1 alleles, enabling fine-tuned modulation of flowering time and explaining domestication-driven shifts from obligate winter to facultative spring types, as documented in varieties from northeastern Pakistan and northwestern India as of 2015 analyses.^[6] Such allelic variation underscores the genetic basis for Pooideae grass adaptations, where vernalization responsiveness traces to an early evolutionary origin within the subfamily.^[23] Legumes, including species like chickpea (Cicer arietinum) and lupin (Lupinus spp.), diverge markedly by lacking functional FLOWERING LOCUS C (FLC) orthologs, the central repressor in Brassicaceae vernalization pathways.^[58] Instead, multiple FT-like paralogs integrate cold-duration signals directly, with vernalization primarily derepressing these florigen genes to trigger reproductive transition without an FLC-mediated repression lift.^[59] This FT-centric mechanism reflects broader phylogenetic divergence, as empirical studies confirm FT upregulation post-cold in legumes correlates with flowering acceleration, distinct from the VRN-dominated repression-induction dynamics in Pooideae cereals.^[60]

Reversal and Modulation

Devernalization by Heat

Devernalization by heat involves the reversal of vernalization-induced flowering competence through exposure to elevated temperatures, which destabilizes the epigenetic silencing of floral repressors. In Arabidopsis thaliana, this process erases the cold-established memory, restoring high levels of the repressor FLOWERING LOCUS C (FLC) expression and delaying bolting.^[61]^[3] Experiments demonstrate that vernalized seedlings exposed to 30°C for 6–7 days immediately after cold treatment exhibit reduced trimethylation of histone H3 at lysine 27 (H3K27me3) at the FLC locus, leading to FLC reactivation, increased vegetative rosette leaf number (e.g., 4 additional leaves), and delayed flowering (e.g., bolting postponed by approximately 5 days compared to stabilized vernalized controls).^[61]^[3] This heat threshold of around 30°C acts within a narrow window; prior stabilization at 20°C for 2 weeks preserves H3K27me3 marks, rendering subsequent 30°C exposure ineffective in reactivating FLC.^[61] The causal mechanism centers on heat-mediated disruption of H3K27me3 stability at FLC, involving partial demethylation that prevents the full propagation of repressive marks required for stable silencing, distinct from the initial cold-dependent nucleation and spreading phases of vernalization.^[61]^[3] In cereals like wheat and rye, heat devernalization nullifies incomplete vernalization competence, as observed in mid-20th-century experiments where warm interruptions (e.g., above 20°C) following short cold periods fully reversed the process, whereas prolonged prior chilling reduced reversibility.^[25] This temperature-sensitive erasure ensures that only sufficiently stable cold memory promotes flowering, adapting plants to variable overwintering conditions.^[25]

Interruptions and Reinitiation of Vernalization

In temperate grasses such as Brachypodium distachyon, brief interruptions by warm temperatures during the vernalization cold period temporarily upregulate floral repressors like VRT2, delaying the stable epigenetic silencing required for flowering competence, but do not fully reset the process. Upon return to cold conditions, vernalization reinitiates through the action of the MADS-box transcription factor BdVRT2, which facilitates renewed repression of flowering inhibitors, preserving partial progress from prior cold exposure. This mechanism ensures that accumulated cold signaling is not entirely lost, differing from complete devernalization where post-vernalization heat fully reactivates repressors and erases stable modifications.^[62] Experimental evidence from B. distachyon accessions shows that interrupting 8 weeks of cold (at 4°C) with 1-2 weeks of warm (22°C) extends the time to flowering by 2-4 weeks compared to uninterrupted controls, even when total cold duration is matched, indicating that the process requires consecutive cold to efficiently propagate silencing. BdVRT2 mutants exhibit heightened sensitivity to these interruptions, failing to reinitiate repression effectively and resulting in prolonged vegetative growth, underscoring its role in bridging interrupted cold phases. Similar patterns occur in related cereals like winter rye, where disruptions delay bolting despite equivalent cumulative cold days, suggesting a conserved quantitative yet non-strictly additive response mediated by VRT2 orthologs.^[62]^[63] Vernalization models incorporating cumulative cold hours—typically calculated as hours below 10°C above a base of 0°C—predict competence thresholds (e.g., 700-1400 hours in wheat relatives), but must account for interruptions reducing efficiency by 20-50% per event in controlled trials. Field validations in temperate cereals confirm that partial credits from interrupted winters correlate with delayed but eventual flowering, as plants reaccumulate units without starting de novo, aiding predictions for variable climates. These dynamics highlight vernalization as a resilient, incremental pathway where epigenetic memory partially endures warm perturbations, enabling adaptation to fluctuating overwinter temperatures.^[62]^[64]

Ecological and Agricultural Roles

Evolutionary Adaptations in Temperate Plants

Vernalization evolved as an adaptive mechanism in temperate plants to synchronize flowering with reliable spring conditions, mitigating risks from fluctuating winter temperatures and false springs that could trigger untimely reproduction. Phylogenetic analyses indicate that vernalization responsiveness originated early in the Pooideae subfamily of grasses, with responsive species distributed widely across the lineage, predating diversification into modern temperate niches.^[23] This trait likely emerged around 40-50 million years ago, coinciding with Eocene-Oligocene cooling and the expansion of seasonal climates, enabling Pooideae ancestors—originally from tropical origins—to colonize cooler latitudes by preventing flowering during brief warm spells amid variable cold periods.^[65] By requiring prolonged non-freezing cold exposure, vernalization ensures reproductive efforts align with post-winter stability, reducing frost damage to developing seeds and enhancing overall fitness in unpredictable environments.^[2] Empirical studies of rear-edge populations, such as those in Campanula americana, reveal ongoing evolutionary adjustments to shorter, less reliable winters, with southern populations exhibiting phenological shifts like reduced vernalization sensitivity or earlier flowering cues to compensate for inconsistent cold durations.^[66] These adaptations highlight vernalization's role in buffering against climate variability, where populations at range margins evolve modified requirements to maintain reproductive timing despite milder conditions, as documented in 2025 analyses comparing rear-edge and central-range genotypes.^[67] Such plasticity underscores the trait's selective advantage in historically glacial-influenced regions, where false thaws posed recurrent threats, allowing plants to defer flowering until cumulative cold verifies seasonal transition.^[66] While vernalization promotes higher seed set by avoiding reproductive losses from late frosts following premature blooms, it imposes constraints in warming scenarios, potentially restricting species distributions if prolonged cold becomes scarce.^[9] In temperate ecosystems, this mechanism has facilitated post-glacial recolonization by favoring genotypes that reliably flower after verified winter passage, though phylogenetic evidence suggests independent evolution in non-Pooideae lineages, indicating convergent selection for cold-cued phenology across angiosperms adapted to seasonal variability.^[68] These adaptations prioritize causal reliability—verifiable cold accumulation—over speculative environmental signals, optimizing survival in climates prone to intermittency.^[2]

Practical Applications in Crop Breeding and Production

Speed vernalization protocols, developed in 2022 for winter cereals including wheat, enable accelerated generation advancement by condensing the required cold exposure period from 6–9 weeks to as little as 2–4 weeks under controlled conditions of 4–10°C with extended photoperiods, facilitating rapid trait stacking and selection for yield-enhancing genes in breeding programs.^[69]^[70] These methods integrate with speed breeding techniques to shorten full generation cycles, allowing breeders to advance winter wheat populations multiple times per year and optimize traits like disease resistance and grain quality without compromising vernalization efficacy.^[71] In practice, such protocols have been applied to winter barley and wheat lines, yielding viable flowering responses and supporting yield improvements through pyramiding of quantitative trait loci for higher productivity.^[72] In field production, vernalization via soil-transmitted cold enables late-emerged winter wheat to fulfill chilling requirements post-emergence, influencing tillering and head development critical to yield potential; Pennsylvania State University extension data from 2025 indicate that plants emerging after mid-November can still achieve partial vernalization if exposed to sufficient sub-4°C soil temperatures over winter, supporting yields of 50–70 heads per square foot with 25–30 plants per square foot under optimal management.^[10] However, incomplete vernalization in such scenarios delays maturity and reduces spike fertility, capping yield at 60–80% of timely-planted counterparts unless compensated by higher plant densities or nitrogen inputs.^[10] Facultative wheat varieties, which exhibit partial vernalization responsiveness, provide verifiable benefits for flexible sowing windows, allowing autumn or spring planting without strict cold mandates and enhancing adaptability to variable weather for sustained yields; a 2025 review highlights their role in optimizing nitrogen uptake and frost tolerance across sowing dates, reducing production risks in temperate regions.^[73] Despite these advantages, over-reliance on accelerated vernalization in breeding risks maladaptation by prioritizing lab-optimized cycles over field-variable cold exposures, potentially yielding cultivars sensitive to heat stress post-flowering or inconsistent chilling, as evidenced by reduced grain yields in vernalized spring wheat under combined abiotic pressures.^[74] Balanced integration with multi-environment trials mitigates such concerns, ensuring robust performance in diverse agroecosystems.^[75]

Recent Advances and Challenges

Discoveries in Epigenetic Dynamics (2020-2025)

In 2022, research identified the histone demethylases JMJ30 and JMJ32 as key regulators that modulate the speed of vernalization in Arabidopsis thaliana by activating the floral repressor FLOWERING LOCUS C (FLC), thereby slowing the accumulation of repressive H3K27me3 marks during cold exposure.^[76] These enzymes counteract premature silencing of FLC, ensuring vernalization requires sufficient cold duration for stable epigenetic repression.^[77] By 2023, studies clarified the dynamic role of the antisense transcript COOLAIR in FLC regulation, showing its induction by C-repeat binding factors (CBFs) during early vernalization stages in Arabidopsis, followed by declining levels as FLC chromatin compacts into a silenced state.^[78] Parallel pathways emerged, with COOLAIR transcription mediating antisense repression independently of Polycomb Repressive Complex 2 (PRC2), which deposits H3K27me3; mutants lacking COOLAIR still achieved partial FLC silencing via PRC2, indicating redundancy in epigenetic silencing mechanisms.^[79] In cereals, epigenomic sequencing advances by 2024 mapped vernalization-responsive cis-regulatory elements at VRN1 loci in wheat, revealing mutations that alter promoter accessibility and cold-induced expression, aiding dissection of quantitative variations in flowering competence.^[80] Extending to legumes, a 2025 analysis of chickpea demonstrated that vernalization upregulates FTa1 and FTa3 expression post-cold, regardless of species-specific flowering responses, linking epigenetic derepression of FLOWERING LOCUS T homologs to accelerated floral transition in this non-model crop.^[81] Also in 2025, screening over 16,000 compounds identified five small-molecule devernalizers (DVRs) that reverse vernalization in Arabidopsis without heat, by reactivating FLC through disruption of H3K27me3 stability and nucleosome positioning at the locus, offering a chemical tool to probe and manipulate epigenetic memory in lab settings.^[3] Concurrently, cold-triggered nucleosome remodeling at FLC was detailed, showing phased chromatin opening and phase separation of PRC2 components as prerequisites for stable silencing, integrating temperature sensing into epigenetic switching.^[48]

Implications for Climate Adaptation and Criticisms of Models

Climate warming reduces the duration and intensity of winter cold periods, potentially providing insufficient vernalization to satisfy requirements in obligate winter cereals, leading to delayed flowering and desynchronization with optimal growth conditions or pollinators.^[82] Field studies in regions like Turkey indicate that many winter wheat genotypes fail to accumulate adequate vernalizing temperatures under projected warming scenarios, compromising adaptation without targeted breeding.^[83] Facultative wheat varieties, with reduced vernalization needs allowing flexible autumn or spring sowing, offer a pathway for resilience to variable winters, as demonstrated in 2025 analyses of their response to climate variability.^[73] However, these adaptations incur yield trade-offs, including lower cold tolerance and altered tillering patterns that diminish biomass accumulation compared to fully vernalized winter types.^[84] Criticisms of predictive models highlight their frequent reliance on Arabidopsis thaliana mechanisms, such as FLC repression, which undervalue the allelic diversity at cereal VRN1, VRN2, and VRN3 loci and the distinct MADS-box gene networks in polyploid species like wheat.^[85] ^[86] This Arabidopsis-centric approach often overestimates adaptive plasticity, ignoring how VRN allelic combinations in cereals modulate responses to fragmented cold exposure more variably than simulated.^[87] Transcriptome studies from 2021 further reveal that vernalization pathways operate independently of cold acclimation stress responses, debunking assumptions in some models that link the two processes and thereby inflating predictions of cross-tolerance under warming.^[88] Empirical field data, such as phenological shifts at species' rear edges, underscore discrepancies with simulations by showing inconsistent compensation for reduced vernalization cues, prioritizing on-ground observations over theoretical projections.^[66] For adaptation strategies, conventional selection from diverse VRN alleles enables incremental gains in vernalization sensitivity tailored to local climates, leveraging standing genetic variation without introducing novel risks, though it proceeds slowly relative to warming rates.^[75] ^[86] Genetic engineering, such as editing VRN loci for reduced requirements, could accelerate targeted adjustments but faces challenges like unintended pleiotropic effects on yield stability and regulatory hurdles, contrasting with selection's preservation of adaptive diversity.^[89] Prioritizing field-validated hybrids over engineered monocultures mitigates uniformity risks in variable environments.^[90]

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Evidence for an Early Origin of Vernalization Responsiveness in ...
Our results demonstrate that vernalization responsive Pooideae species are widespread, suggesting that this trait evolved early in the lineage.Missing: discovery | Show results with:discovery
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Historic rewiring of grass flowering time pathways and implications ...
Dec 26, 2024 · Many temperate grasses use vernalization as a cue to ready their development for spring flowering. ... Evidence for an early origin of ...
[25]
VERNALIZATION AND ITS RELATIONS TO DORMANCY1
The very notion of vernalization originated from this early agricultural experience.
[26]
Preliminary experiments on vernalisation
Mar 27, 2009 · 2. In addition to early developmental stimulation, winter varieties of barley and wheat also showed marked acceleration in the date of earing.
[27]
Vernalization - an overview | ScienceDirect Topics
It was Gassner who, between 1910 and 1918, studied the floral response to low temperatures, and introduced the concept of the requirement of a cold treatment ...
[28]
Vernalization in cereals - PMC - PubMed Central
Jun 22, 2009 · Plant breeders have known for many decades that some crops require vernalization. Examples of these are the winter wheats and winter barleys ...
[29]
Lysenkoism Against Genetics: The Meeting of the Lenin All-Union ...
Lysenko believed that heritable changes arise in plants as a result of vernalization, while geneticists already knew the idea to be false. The Ukrainian ...
[30]
Russia's new Lysenkoism - ScienceDirect.com
Oct 9, 2017 · Lysenkoism was a neo-Lamarckian idea, claiming that in crop plants, such as wheat, environmental influences are heritable via all cells of the organism.
[31]
Science and politics - PMC - NIH
Lysenko's rejection of Mendelian genetics in favour of Lamarckism, his embrace of the hybridization theories of Russian horticulturist Ivan Michurin (1855–1935 ...
[32]
The Lysenko effect: undermining the autonomy of science
The Lysenko effect was the suppression of genetics in Soviet Russia, showing how politics can corrupt science, and how indirect political interference can ...
[33]
Inherit a Problem: How Lysenkoism Ruined Soviet Plant Genetics ...
Jan 20, 2020 · Lysenkoism, supported by Stalin, ruined Soviet agriculture and genetics, leading to famine and the inability to feed the people, and was a ...
[34]
The pushback against state interference in science - PubMed Central
Nov 5, 2021 · In 1948, Konstantinov wrote a letter to Stalin analyzing the damage caused by many of Lysenko's gambles: wide-scale vernalization, direct ...
[35]
Lysenko and Russian genetics: an alternative view - Nature
Jul 26, 2017 · For example, it was Lysenko who coined the term vernalization, which is now still an extant scientific term and frequently appears in Nature ...Missing: impact | Show results with:impact
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Lysenkoism as an accusatory term for state-led, typically Communist or (more recently) left wing, manipulation of science for political ends.<|separator|>
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A PHD-Polycomb Repressive Complex 2 triggers the epigenetic ...
Nov 4, 2008 · In summary, we have defined a sequence of events required for the vernalization-induced repression of FLC by a Polycomb mechanism (Fig. 4).
[38]
Antisense COOLAIR mediates the coordinated switching of ... - PNAS
Vernalization, the quantitative acceleration of flowering by prolonged cold, leads to the epigenetic silencing of FLC (1, 3). Polycomb Repressive Complex 2 ( ...<|separator|>
[39]
Vernalization and Epigenetic Inheritance: A Game of Histones
Dec 18, 2017 · The Polycomb repressive complex 2 (PRC2) modifies chromatin by catalyzing trimethylation of lysine 27 on histone H3 (H3K27me3), a mark required ...
[40]
Vernalization – a cold-induced epigenetic switch
Aug 15, 2012 · The basis of mitotic stability of histone modifications. Any long-term epigenetic memory must be maintained through multiple rounds of cell ...Introduction · Epigenetic Silencing After... · Generic Principles That Have...
[41]
Quantitative effects of vernalization on FLC and SOC1 expression
We present results that provide an insight into the quantitative regulation of FLC by vernalization. Analysis of the effect of seed or plant cold treatment on ...
[42]
Targets and mechanisms of epigenetic regulation in the temperate ...
Aug 4, 2025 · Vernalisation is a prolonged cold exposure that synchronises flowering with favourable seasonal conditions, protecting reproductive ...
[43]
Vernalization treatment induces site-specific DNA hypermethylation ...
Vernalization requires epigenetic silencing of FLC by histone methylation. ... Kismeth: Analyzer of plant methylation states through bisulfite sequencing.
[44]
Duration of cold exposure defines the rate of reactivation of ... - Nature
Sep 29, 2020 · The duration of cold defines the rate of AhgFLC reactivation via the buffering function of H3K27me3 against temperature increase.
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Control of Arabidopsis flowering: the chill before the bloom
Aug 15, 2004 · FLC is a MADS-box transcriptional repressor, expressed predominantly in shoot and root apices and vasculature, that quantitatively represses ...The Floral Repressor, Flc · Activation Of Flc · Hdacs In The Flowering...
[46]
The Role of VIN3-LIKE Genes in Environmentally Induced ... - NIH
VIN3-LIKE 1 (VIL1) is involved in mediating the flowering response to both cold and day length via regulation of two related genes.
[47]
Vernalization requires epigenetic silencing of FLC by histone ...
Jan 8, 2004 · VRN1 encodes a DNA-binding protein whereas VRN2 encodes a homologue of one of the Polycomb group proteins, which maintain the silencing of genes ...
[48]
Cold-induced nucleosome dynamics linked to silencing of ... - Nature
Jul 1, 2025 · The vrn1 mutation compromises FLC silencing, as previously observed in a different Arabidopsis genotype. Moreover, the effect of vrn1 on FLC ...
[49]
Genetic and Epigenetic Mechanisms Underlying Vernalization
Jan 1, 2014 · Mutations in autonomous pathway genes result in high level of FLC expression and thus confer the vernalization requirement for early flowering ...
[50]
FCA, a Gene Controlling Flowering Time in Arabidopsis, Encodes a ...
The balance of the autonomous and vernalization-dependent promotion pathways may explain the different flowering times and vernalization responses of the ...
[51]
Loss of FLOWERING LOCUS C Activity Eliminates the Late ...
Under LD conditions, the late-flowering phenotype of autonomous pathway mutations (fca, fpa, fve, and ld) was completely eliminated in the presence of a null ...
[52]
Autonomous Pathway: FLOWERING LOCUS C Repression through ...
Martin Koornneef and others in the 1990s and 2000s identified fpa, fca, fy, fve, flk, ld, and fld as mutants of the autonomous pathway (Lee et al., 1994; Sanda ...
[53]
Competence to Flower: Age-Controlled Sensitivity to Environmental ...
Age-related responsiveness to vernalization controlled by the miR156/SPL and miR172/AP2 · Spatially distinct roles of SPL genes in the Arabidopsis SAM.
[54]
Molecular basis of age-dependent vernalization in Cardamine ...
May 31, 2013 · We demonstrate that the levels of two age-regulated microRNAs, miR156 and miR172, regulate the timing of sensitivity in response to vernalization.
[55]
Regulation of flowering time by the miR156-mediated age pathway
Jun 22, 2014 · Although the role of miR156 and miR172 in setting a threshold for the sensitivity in response to vernalization is conserved between A. alpina ...
[56]
Epigenomic identification of vernalization cis-regulatory elements in ...
Jul 30, 2024 · Our study offers critical insights into wheat vernalization's epigenomic dynamics and identifies the putative regulatory elements crucial for developing wheat ...Missing: reviewed | Show results with:reviewed<|separator|>
[57]
The wheat and barley vernalization gene VRN3 is an orthologue of FT
VRN3 is a promoter of flowering up-regulated by LD, which positively regulates VRN1, the meristem identity gene.
[58]
Mechanisms of Vernalization-Induced Flowering in Legumes - PMC
Aug 31, 2022 · Despite the absence of FLC orthologs, the legume FT genes retain the role of major vernalization signal integrators (Table 2). Legumes have ...
[59]
Modeling the Flowering Activation Motif during Vernalization in ...
Dec 23, 2023 · The orthologs of the FLC gene, a major regulator of vernalization response in Arabidopsis, are absent or non-functional in the vernalization- ...
[60]
Photoperiod and Vernalization Control of Flowering-Related Genes
Oct 25, 2020 · Lupins, like many other legumes, do not have any FLC homologs. ... However, in many legumes, FT genes were found to be associated with flowering ...
[61]
Heat can erase epigenetic marks of vernalization in Arabidopsis - PMC
We reported that high temperature leads to devernalization and, consistently, to FLC reactivation in Arabidopsis seedlings. Here we show that the repressive ...
[62]
Transcription factor VRT2 reinitiates vernalization when interrupted ...
Sep 24, 2024 · Despite these independent origins, it appears that factors regulating vernalization are commonly targeted during devernalization. In Arabidopsis ...
[63]
[PDF] Transcription factor VRT2 reinitiates vernalization when ... - Lirias
Previous reports in winter rye have shown that disrupting the vernaliza- tion process can delay flowering time, despite the total sum of vernalizing days being ...
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A vernalization-intensity model to predict bolting in sugar beet
Aug 7, 2025 · They expressed the length and intensity of cold treatment as vernalization-weighted hours (vwh) and modeled the bolting rate of sugar beet in ...
[65]
Successive evolutionary steps drove Pooideae grasses from tropical ...
Nov 1, 2017 · Evidence for an early origin of vernalization responsiveness in temperate Pooideae grasses. Plant Physiology 172: 416–426. 10.1104/pp ...
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Shifts in vernalization and phenology at the rear edge hold insight ...
Mar 6, 2025 · Temperate plants often regulate reproduction through winter cues, such as vernalization, that may decrease under climate change.
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Shifts in vernalization and phenology at the rear edge hold insight ...
We studied how rear-edge populations have adapted to shorter winters and compared them to the rest of the range in the herb Campanula americana. Using ...
[68]
Independent recruitment of FRUITFULL-like transcription factors in ...
Our results demonstrate that vernalization-induced flowering has evolved multiple times independently in at least five grass subfamilies.
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Speed vernalization to accelerate generation advance in winter ...
Aug 1, 2022 · Vernalization is the requirement for a prolonged period of cold before certain plants can transition from vegetative to reproductive development ...
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Speed vernalization to accelerate generation advance in winter ...
Aug 1, 2022 · The speed vernalization protocol developed in this study has great potential to accelerate biological research and breeding outcomes for winter crops.
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Acceleration of wheat breeding: enhancing efficiency and practical ...
Nov 4, 2023 · One is the vernalization room, which is kept at 8–10 °C with a photoperiod of 22 h of light [9]. These vernalization conditions must be ...
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[PDF] A new protocol for speed vernalisation of winter cereals - bioRxiv
Dec 6, 2021 · Vernalisation is the requirement for a prolonged period of cold before certain plants can. 29 transition from vegetative to reproductive ...
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https://pubs.acs.org/doi/full/10.1021/acsagscitech.5c00039
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(PDF) Impact of Vernalization and Heat Stress on Spring Wheat ...
Sep 6, 2024 · Vernalization significantly decreased grain yield of spring wheat. Under non-vernalization conditions, heat stress after flowering produced a ...
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Crop adaptation to climate change as a consequence of long-term ...
Nov 22, 2020 · Long-term genetic gain through conventional breeding has incrementally increased yields of modern crops through accumulation of beneficial, small-effect ...
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Histone Demethylases JMJ30 and JMJ32 Modulate the Speed of ...
Jun 30, 2022 · Devernalization is a reversion of vernalized status to non-vernalized status by heat. It was reported that H3K27me3 on the FLC locus was reduced ...
[77]
Histone Demethylases JMJ30 and JMJ32 Modulate the Speed of ...
Jul 1, 2022 · These results suggested that the histone demethylases JMJ30 and JMJ32 brake vernalization through the activation of FLC.
[78]
Vernalization-triggered expression of the antisense transcript ... - eLife
Feb 1, 2023 · Subsequently, upon returning to warm temperatures, H3K27me3 marks are spread over the entire FLC chromatin region by LIKE HETEROCHROMATIN ...<|control11|><|separator|>
[79]
COOLAIR and PRC2 function in parallel to silence FLC ... - PNAS
We show that COOLAIR transcription and PRC2 (Polycomb Repressive Complex 2) silence FLC in parallel pathways: an antisense-mediated transcriptional repression.
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Epigenomic identification of vernalization cis-regulatory elements in ...
Jul 30, 2024 · Sequencing 4.3 million mutations in wheat promoters to understand and modify gene expression. Proc Natl Acad Sci USA. 2023;120(38):e2306494120.Missing: modeling interruptions
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Effect of Vernalization on the Expression of Flowering Time Genes in ...
The expression levels of FTa1 and FTa3 gene products increased significantly after vernalization, regardless of species. At the same time, the response to ...
[82]
Vernalization requirement duration in winter wheat is controlled by ...
Winter wheat requires a period of low temperatures to accelerate flowering (vernalization). This requirement could make winter wheat more vulnerable to ...
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Winter Wheat Adaptation to Climate Change in Turkey - MDPI
Our research findings showed that many winter wheat genotypes would not successfully adapt to climate change.<|separator|>
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Yield and yield components of facultative wheat are affected by ...
It exhibits lower cold tolerance and lower vernalization requirement compared to autumn-sown wheat, which allows spring-sowing and autumn-sowing with mild ...
[85]
Targets and mechanisms of epigenetic regulation in the temperate ...
Aug 5, 2025 · The goal of this review is to synthesise recent advances in our understanding of the epigenetic regulation of vernalisation in temperate cereals ...
[86]
Allelic Variation at the Vernalization Response (Vrn-1 ... - Frontiers
Jun 22, 2020 · Models for the genetic control of flowering in cereals suggest a functional relationship between the vernalization- and photoperiod-response ...
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The molecular basis of vernalization-induced flowering in cereals
A comparison of flowering time pathways in cereals and Arabidopsis shows that the vernalization response is controlled by different MADS box genes, but ...<|separator|>
[88]
Transcriptome analysis reveals Vernalization is independent of cold ...
Time-lag initiation of the two pathways and weighted gene co-expression network analysis (WGCNA) revealed that vernalization is independent of cold acclimation.Missing: debunked | Show results with:debunked
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[PDF] Selection constraints of plant adaptation can be relaxed by gene ...
Jun 11, 2024 · Future climates are predicted to escalate conflicts in natural selection among adaptive traits, but our work shows that surprisingly simple ...
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Plant breeding for climate change: Opportunities for adaptation and ...
Plant breeding for climate change uses diverse genetic resources to adapt crops to climate change and breed plants that can mitigate those conditions.