Biology subjects

Hytonen, T.

Publications and source records attributed to Hytonen, T..

10 recordsLinked to original sources

bioRxiv2026

TFL1 homolog integrates temperature signals to control timing of developmental transitions in woodland strawberry

In overwintering plants of temperate climates, inflorescence and flower buds commonly form in autumn under cool temperatures, while flowering occurs the following spring. However, how cool temperatures regulate inflorescence and flower development remains poorly understood. We used woodland strawberry (Fragaria vesca L.), a perennial Rosaceae herb, to investigate the effects of cool temperature on reproductive development. We characterized temperature-dependent developmental patterns at macroscopic and microscopic scale and combined time- and stage-specific gene expression analyses. We show that cool temperature accelerates floral transition but subsequently slows inflorescence and flower development, causing increased inflorescence branching, higher floral organ numbers, and berry fasciation. These effects were restricted to the apical inflorescence and flower meristems, whereas higher-order meristems remained unaffected. Expression analyses revealed stage- and temperature-dependent changes in the abundance and spatial distribution of TERMINAL FLOWER1 (FvTFL1) encoding the floral repressor. Loss of functional FvTFL1 abolished temperature-dependent differences in floral transition, inflorescence branching, and floral organ numbers. Our results establish FvTFL1 as a major regulator of temperature-responsive reproductive development. The stage-specific activity of FvTFL1 provides a mechanism through which woodland strawberry integrates temperature signals to balance developmental plasticity of the primary growth axis with the robustness of lateral branches.

plant biology↗

bioRxiv2026

FvTFL1 reverses the function of FvGI-FvCO-FvFT1 pathway in the photoperiodic flowering of woodland strawberry

O_LIPhotoperiod is a stable seasonal signal. Although the photoperiodic flowering is well understood in short-day (SD) and long-day (LD) annual plants, regulatory mechanisms in perennials remain elusive. In a perennial woodland strawberry (Fragaria vesca L.), flowering is induced in SDs in autumn and plants flower following spring, while in plants with mutated FvTERMINAL FLOWER1 (FvTFL1), LDs induce flowering. C_LIO_LIWe investigated photoperiodic flowering of F. vesca through phenotypic and molecular characterization of transgenic lines and their crosses. We studied natural variation in flowering time and gene expression in European accessions, and explored their correlations with climatic, geographical and genetic origins. C_LIO_LIWe showed that FvGIGANTEA (FvGI) and FvCONSTANS (FvCO) activate FvFLOWERING LOCUS T1 (FvFT1) in LDs resulting in early flowering in fvtfl1 mutant, while in SD F. vesca, activation of FvTFL1 by FvFT1 reverses the photoperiodic requirement of flowering. In natural accessions, decreasing expression of FvFT1 and FvTFL1 towards colder climates in the east and north correlated with earlier flowering. C_LIO_LIWe define a photoperiodic flowering mechanism controlling floral transition of perennial F. vesca in autumn that differs from known mechanisms in annual and perennial plants. Our findings open new avenues to understand how perennial plants cope with changing seasons across climatic and geographical ranges. C_LI

plant biology↗

bioRxiv2025

FvFT1-FvTFL1 epistasis drives flowering time adaptation in woodland strawberry

Floral transition and flower bud development during autumn and winter are common in plants inhabiting temperate and cold climates. However, our understanding of the natural variation and underlying molecular mechanisms of such life-history strategy remain largely limited to Brassicaceae model species. To address this gap, we investigated flowering time variation across a broad geographical and bioclimatic range in Fragaria vesca (woodland strawberry), a perennial herb from the Rosaceae family. We found that the timing of floral transition and subsequent flowering in F. vesca populations is primarily shaped by population structure, reflecting postglacial colonization patterns from distinct refugia, and is further modulated by adaptations to winter temperatures. The association of increasing expression of FvFT1, the strawberry ortholog of FLOWERING LOCUS T, with delayed flowering in accessions from diverse climatic and genetic backgrounds suggested FvFT1 as a major candidate gene regulating flowering time variation. We further demonstrated how distinct florigen and antiflorigen roles of FvFT1 are dependent on functional alleles of F. vesca TERMINAL FLOWER1 (FvTFL1) and how the epistatic relationship between FvFT1 and FvTFL1 controls adaptive flowering time variation by adjusting sensitivity to floral inductive photoperiod and vernalization. Our findings reveal a major molecular basis of how timing of floral transition has adapted to temperature in a Rosaceae model inhabiting multiple climatic zones and highlight lineage-specific adaptive mechanisms underlying convergent life-history strategy. Significance statementMolecular mechanisms underlying flowering time adaptation have been primarily investigated in annual plants that transit to flowering in spring. However, many perennial plant species in temperate climates form flower buds in autumn and winter to flower in the following spring. We uncover wide geographical and climatic patterns of flowering time adaptation in the perennial model species woodland strawberry. We report that natural variation in the expression of strawberry FLOWERING LOCUS T (FvFT1) gene originating ice ages ago and further driven by local temperature adaptations control flowering time variation in this species. Finally, we show how FvFT1 may function as both flowering promoter and repressor depending on the genetic background, thus revealing a novel mechanism of flowering time adaptation.

plant biology↗

bioRxiv2025

Naturally occurring variation in gene-associated transposable elements impacts gene expression and phenotypic diversity in woodland strawberry

Transposable elements (TEs) are major components of plant genomes and powerful drivers of regulatory and phenotypic diversity, yet their mechanistic contributions to gene regulation in non-model crops remain poorly understood. Here, we generate a comprehensive and curated TE annotation for woodland strawberry (Fragaria vesca), revealing that TEs comprise [~]30% of the genome and uncovering an exceptional enrichment of gene-proximal miniature inverted-repeat transposable elements (MITEs) and short inverted repeats (IRs). Integrating small RNA, DNA methylation and transcriptomic data across tissues and fruit developmental stages, we show that most gene-associated MITEs and IRs are epigenetically active, producing 24-nt siRNAs and displaying dynamic CHH methylation patterns during fruit ripening. Genome-wide analyses reveal that MITE- and IR-proximal genes occupy a distinct regulatory landscape compared to genes associated with other TE classes, consistent with a role in context-dependent transcriptional regulation. As a mechanistic proof of concept, we demonstrate that a Mutator-like MITE located near FvAAT1, encoding a key alcohol acyltransferase required for aroma ester biosynthesis, modulates gene expression through siRNA-dependent DNA methylation and ripening-stage-specific chromatin reorganization. Transient perturbation of this MITE epigenetic state alters FvAAT1 expression, and chromatin conformation capture reveals MITE-associated looping at this locus. Notably, this regulatory MITE is conserved in commercial strawberry species, suggesting an evolutionarily preserved function in fruit quality traits. Extending these findings to natural variation, we characterize thousands of TE insertion and deletion polymorphisms across 210 European F. vesca accessions and show that TE-based variation recapitulates population structure. A TE-informed genome-wide association analysis identifies significant associations between MITE and IR polymorphisms and fruit volatile compounds, including key aroma determinants. Together, our results establish MITEs and short IRs as widespread, epigenetically active regulatory elements that shape gene expression, chromatin architecture and phenotypic diversity in strawberry. This work provides a conceptual and methodological framework to integrate TE-driven regulatory variation into population genomics, breeding and genome-editing strategies aimed at improving fruit quality.

genomics↗

bioRxiv2024

Late Quaternary climatic impact on the woodland strawberry genome: a perennial herb's tale

Exploring a species paleohistory is crucial for understanding its responsiveness to climatic events, identifying drivers of adaptation, and developing effective biodiversity conservation strategies in the face of ongoing climate change. We analyzed 200 genomes of the perennial herb woodland strawberry (Fragaria vesca L.) from across Europe and investigated the population structure and demographic history of the species during past geoclimatic events. We found a clear division of populations into western and eastern genetic clusters, indicative of distinct glacial refugia and adaptations to variation in temperature seasonality. The eastern core populations were several times larger (defined as effective population size, NE) than populations in other regions, showed no evidence of inbreeding, and were resilient to several glacial maxima. However, we observed decreasing NE and higher inbreeding in populations toward range edges, particularly in the north, where these individuals went through bottlenecks during glaciations. Population divergence suggested that western and eastern Europe were colonized from separate refugia in multiple waves during the Holocene, while the largest current populations from the northern Mediterranean to southern regions of the Nordic countries formed a connected population chain with gene flow between eastern core populations and western Europe, primarily occurring through Central Europe. Similar patterns of colonization and hybridization may have occurred during past interglacial periods, contributing to the present-day population structure of woodland strawberry. We suggest that the unprecedented resolution of the species climatic history across six glacial-interglacial cycles presented here holds the promise of transforming the general understanding of species paleohistory through geoclimatically tracing ancestral haplotypes.

plant biology↗

bioRxiv2022

Diversity of the woodland strawberry inflorescences results from heterochrony antagonistically regulated by FvTFL1 and FvFT1

A vast variety of inflorescence architectures have formed during angiosperm evolution. Here we analyze the diversity and development of the woodland strawberry inflorescence. We show that it is a thyrse: a compound inflorescence in which the primary monopodial axis supports lateral sympodial branches, thus combining features of racemes and cymes. We demonstrate that this architecture is related to differences in the size and shape of the primary and lateral inflorescence meristems. We further show that woodland strawberry homologs of TERMINAL FLOWER 1 (TFL1) and FLOWERING LOCUS T (FT) antagonistically regulate the development of both the racemose and cymose components of the strawberry thyrse: the loss of functional FvTFL1 and overexpression of FvFT1 reduce the number and complexity of the cymose components, whereas silencing of FvFT1 has the opposite effect and can partially rescue the fvtfl1 mutation. We complement our experimental findings with a computational model, which captures the development of the woodland strawberry inflorescence using a small set of rules, and shows that its phenotypic diversity can be explained in terms of heterochrony resulting from the opposite action of FvTFL1 and FvFT1 on the progression from the branching to flowering state.

plant biology↗

bioRxiv2022

Temperature-induced methylome changes during asexual reproduction trigger transcriptomic and phenotypic changes in Fragaria vesca

Plants must quickly adapt to a changing environment in order to maintain their fitness. One rapid adaptation mechanism that promotes plasticity is epigenetic memory, which may provide long-lived organisms the precious time needed to adapt to climate change. In this study, we used the perennial plant Fragaria vesca as a model to determine how the methylome and transcriptome adapt to elevated temperatures (28 vs. 18 {degrees}C) over three asexual generations. Changes in flowering time, stolon number, and petiole length were induced in responses to temperature treatment in one or more ecotypes after three asexual generations in a manner indicative of an epigenetic memory. Induced methylome changes differed between four different ecotypes from Norway, Iceland, Italy, and Spain, but there were also some shared responses. Elevated temperature conditions induced significant phenotypic and methylation changes, particularly in the Norwegian ecotype. Most of the differentially methylated regions (DMRs) were in the CHG context, and most CHG and CHH DMRs were hypermethylated. Notably, the four ecotypes shared only eight CHG DMR peaks. Several differentially methylated genes (DMGs) also showed a change in gene expression. Ecotype-specific methylation and expression patterns were observed for genes related to gibberellin metabolism, flowering time, epigenetics. Furthermore, when repetitive elements (REs) were found near ({+/-}2 kb) or inside a gene, they showed a negative correlation with gene expression. In conclusion, phenotypic changes induced by elevated temperatures during asexual reproduction were accompanied by changes in DNA methylation patterns. Also, positional influences of REs impacted gene expression, indicating that DNA methylation may be involved in both general and ecotype-specific phenotypic plasticity in F. vesca.

genomics↗

bioRxiv2022

Epistatic Modifiers Influence the Expression of Continual Flowering in Strawberry

O_LIPrevious work within the community led to the identification of a single dominant allele that controls the everbearing trait. However, frequent observations have indicated that crosses do not segregate in a Mendelian fashion, as would be expected for a trait controlled by a single dominant gene. Therefore, it was hypothesised that one or more unidentified epistatic alleles interact with the major gene. C_LIO_LIA GWAS was conducted on 587 June-bearers and 207 everbearers to assess the genetic components associated with flowering habit. The segregation ratios of parental strawberry lines with known phenotypes were used to validate the identified alleles. C_LIO_LIThree loci including the known, major FaPFRU locus and two epistatic modifiers were identified. These modifiers function as enhancers of the everbearing trait in individuals containing a single copy of the FaPFRU everbearing allele and appear to be functionally redundant. Principally, heterozygous individuals required the presence of two modifying alleles in order to allow expression of the everbearing trait. C_LIO_LIInclusion of the epistatic alleles improved the prediction of everbearing segregation ratios, beyond that of a single allele model, however, a large proportion of the variation remained unexplained. Future work should identify the additional repressor and enhancer modifiers not identified here. Discovering the genetic components controlling the everbearing trait can enable genetic informed strawberry improvement. C_LI Society Impact StatementUntil the 1970s, the majority of commercial strawberry varieties produced a single bloom of flowers. However, continuously flowering, everbearing strawberries are now routinely cultivated and use is increasing. Indeed, introgression of the everbearing flowering trait can lead to economic benefits for growers through the production of a continual crop from the same plant. Utilising genetic guided improvement has the power to streamline everbearing generation. As such, utilising the genetic markers reported here, can help to identify everbearing individuals at an early time point in the breeding process. Furthermore, these markers can help to improve the predictions of progeny segregation ratios.

plant biology↗

bioRxiv2021

Natural variation in the control of flowering and shoot architecture in diploid Fragaria species

In perennial fruit and berry crops of the Rosaceae family, flower initiation occurs in late summer or autumn after downregulation of a strong repressor TERMINAL FLOWER1 (TFL1) and flowering and fruiting takes place the following growing season. Rosaceous fruit trees typically form two types of axillary shoots, short flower-bearing shoots called spurs and long shoots that are respectively analogous to branch crowns and stolons in strawberry. However, regulation of flowering and shoot architecture differs between species and environmental and endogenous controlling mechanisms have just started to emerge. In woodland strawberry (Fragaria vesca L.), long days maintain vegetative meristems and promote stolon formation by activating TFL1 and GIBBERELLIN 20-OXIDASE4 (GA20ox4), respectively, while silencing of these factors by short days and cool temperatures induces flowering and branch crown formation. We characterized flowering responses of 14 accessions of seven diploid Fragaria species native to diverse habitats in the northern hemisphere, and selected two species with contrasting environmental responses, F. bucharica Losinsk. and F. nilgerrensis Schlecht. ex J. Gay for detailed studies together with F. vesca. Similar to F. vesca, F. bucharica was induced to flower in short days at 18{degrees}C and regardless of photoperiod at 11{degrees}C after silencing of TFL1. F. nilgerrensis maintained higher TFL1 expression level and likely required cooler temperatures or longer exposure to inductive treatments to flower. We also found that high expression of GA20ox4 was associated with stolon formation in all three species, and its downregulation by short days and cool temperature caused branch crown formation in F. vesca and F. nilgerrensis, although the latter did not flower. F. bucharica, in contrast, rarely formed branch crowns, regardless of flowering or GA20ox4 expression level. Our findings highlighted diploid Fragaria species as a rich source of genetic variation controlling flowering and plant architecture, with potential applications in breeding of Rosaceous crops.

plant biology↗

bioRxiv2020

Allelic Variation of MYB10 is the Major Force Controlling Natural Variation of Skin and Flesh Color in Strawberry (Fragaria spp.) fruit

Anthocyanins are the principal color-producing compounds synthesized in developing fruits of strawberry (Fragaria spp.). Substantial natural variation in color have been observed in fruits of diploid and octoploid accessions, resulting from distinct accumulation and distribution of anthocyanins in fruits. Anthocyanin biosynthesis is controlled by a clade of R2R3 MYB transcription factors, among which MYB10 has been shown as the main activator in strawberry fruit. Here, we show that MYB10 mutations cause most of the anthocyanin variation observed in diploid woodland strawberry (F. vesca) and octoploid cultivated strawberry (F. xananassa). Using a mapping-by-sequencing approach, we identified a gypsy-transposon insertion in MYB10 that truncates the protein and knocks out anthocyanin biosynthesis in a white-fruited F. vesca ecotype. Two additional loss-of-function MYB10 mutations were identified among geographically diverse white-fruited F. vesca ecotypes. Genetic and transcriptomic analyses in octoploid Fragaria spp. revealed that FaMYB10-2, one of three MYB10 homoeologs identified, residing in the F. iinumae-derived subgenome, regulates the biosynthesis of anthocyanins in developing fruit. Furthermore, independent mutations in MYB10-2 are the underlying cause of natural variation in fruit skin and flesh color in octoploid strawberry. We identified a CACTA-like transposon (FaEnSpm-2) insertion in the MYB10-2 promoter of red-fleshed accessions that was associated with enhanced expression and anthocyanin accumulation. Our findings suggest that putative cis regulatory elements provided by FaEnSpm-2 are required for high and ectopic MYB10-2 expression and induction of anthocyanin biosynthesis in fruit flesh. We developed MYB10-2 (sub-genome) specific DNA markers for marker-assisted selection that accurately predicted anthocyanin phenotypes in octoploid segregating populations.

plant biology↗