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Dormancy is an adaptation mechanism of plants to environmental stress. Myricaria laxiflora undergoes a long period of flooding stress every year. In order to determine whether this species escapes flooding stress through dormancy, young branches and leaves were collected at different time points before the onset of flooding, and changes in the content/activity of hormones/enzymes that are closely involved in plant growth were monitored. The inducing environmental factors of summer dormancy were identified. The branches and leaves of M. laxiflora showed the following trends as summer flooding approached: (1) gradual increase in the abscisic acid content; (2) gradual decrease in the gibberellin and cytokinin contents; and (3) a continuous decrease in the activities of malate dehydrogenase (MDH), ribulose diphosphate carboxylase (RuBisCo), and glycolate oxidase (GLO). Pearson correlation analysis revealed (1) daylight duration was highly correlated with the hormone content and enzyme activity; (2) the daily mean air temperature (DMAT) was significantly correlated with the cytokinin content. These findings suggest that daylight duration was the main environmental factor leading to changes in the phytohormone content and enzyme activity as well as leading to summer dormancy. M. laxiflora undergoes dormancy before the onset of summer flooding to escape summer flooding stress. Our data indicate that summer flooding does not impede the survival and growth of M. laxiflora.

Repression of embryonic traits during the seed-to-seedling phase transition requires the inactivation of master transcription factors associated with embryogenesis. How the timing of such inactivation is controlled is unclear. Here, we report on a novel transcriptional co-repressor, Arabidopsis thaliana SDR4L, that forms a feedback inhibition loop with the master transcription factors LEC1 and ABI3 to repress embryonic traits post-imbibition. LEC1 and ABI3 regulate their own expression by inducing AtSDR4L during mid to late embryogenesis. AtSDR4L binds to sites upstream of LEC1 and ABI4, and these transcripts are upregulated in Atsdr4l seedlings. Atsdr4l seedlings phenocopy a LEC1 overexpressor. The embryonic traits of Atsdr4l can be partially rescued by impairing LEC1 or ABI3. The penetrance and expressivity of the Atsdr4l phenotypes depend on both developmental and external cues, demonstrating the importance of AtSDR4L in seedling establishment under suboptimal conditions.

Background and Aims Environments experienced by both parents and offspring influence progeny traits, but the epigenetic mechanisms that regulate the balance of parental vs. progeny control of progeny phenotypes are not known. We tested whether DNA methylation in parents and/or progeny mediates responses to environmental cues experienced in both generations. Methods Using Arabidopsis thaliana, we manipulated parental and progeny DNA methylation both chemically, via 5-azacytidine, and genetically, via mutants of methyltransferase genes, then measured progeny germination responses to simulated canopy shade in parental and progeny generations. Key Results We first found that germination of offspring responded to parental but not seed demethylation. We further found that parental demethylation reversed the parental effect of canopy in seeds with low (Cvi-1) to intermediate (Col) dormancy, but it obliterated the parental effect in seeds with high dormancy (Cvi-0). Demethylation did so by either suppressing germination of seeds matured under white-light (Cvi-1) or under canopy (Cvi-0), or by increasing the germination of seeds matured under canopy (Col). Disruption of parental methylation also prevented seeds from responding to their own light environment in one genotype (Cvi-0, most dormant), but it enabled seeds to respond to their own environment in another genotype (Cvi-1, least dormant). Using mutant genotypes, we found that both CG and non-CG DNA methylation were involved in parental effects on seed germination. Conclusions Parental methylation state influences seed germination more strongly than does the progeny’s own methylation state, and it influences how seeds respond to environments of parents and progeny in a genotype-specific manner.

Pre-harvest sprouting (PHS) in cereal crops, is a universal phenomenon that affects grain yield and quality (Tai et al. 2021). Breeders use different methods to strengthen seed dormancy to prevent PHS including genome editing. Abscisic acid is one of the phytohormones that promote seed dormancy. ABA8ox genes encode abscisic acid 8’ hydroxylase (ABA8OX) which determines the level of ABA content by catabolizing ABA (Vallabhaneni and Wurtzel 2010). Three genes in rice encode ABA8’ hydroxylases, namely OsABA8ox1 (LOC_Os02g47470), OsABA8ox2 (LOC_Os08g36860) and OsABA8ox3 (LOC_Os09g28390) (Kushiro et al. 2004; Cai et al. 2015; Zhang et al. 2020). Each gene is expressed in different tissues of rice.

Seeds exhibit primary dormancy to prevent germination under unfavourable conditions. Previous studies have shown that the gibberellin signalling intermediate RGA-LIKE2 (RGL2) forms a transcription factor complex with DNA-BINDING ONE ZINC FINGER6 (DOF6) in regulating seed dormancy in Arabidopsis. Using an RNA-sequencing approach, we identified MAJOR LATEX PROTEIN-LIKE PROTEIN329 (MLP329) as a downstream target of DOF6. MLP329 was found to be a positive regulator of primary seed dormancy, because freshly harvested unstratified mlp329 mutant seeds showed early germination, while unstratified transgenic seeds overexpressing MLP329 showed poor germination. MLP329 expression level was reduced in wild-type seeds upon dry storage and cold stratification. MLP329 expression level was enhanced by DOF6; however, DOF6-dependent MLP329 expression was suppressed in the presence of RGL2. MLP329 expression was enhanced in seeds treated with ABA and auxin IAA. Moreover, the mlp329 mutant seeds exhibited enhanced expression of the GA biosynthetic gene GA1 and suppression of the ABA biosynthetic gene ZEP compared to the overexpression lines. The observed suppression of DOF6-dependent MLP329 expression by RGL2 reveals a possible negative feedback mechanism to modulate seed dormancy. MLP329 also probably enhances the endogenous ABA/GA ratio to positively regulate primary seed dormancy.

Bud dormancy helps woody perennials survive winter and activate robust plant development in the spring. For apple (Malus × domestica), short-term chilling induces bud dormancy in autumn, then prolonged chilling leads to dormancy release and a shift to a quiescent state in winter, with subsequent warm periods promoting bud break in spring. Epigenetic regulation contributes to seasonal responses such as vernalization. However, how histone modifications integrate seasonal cues and internal signals during bud dormancy in woody perennials remains largely unknown. Here, we show that H3K4me3 plays a key role in establishing permissive chromatin states during bud dormancy and bud break in apple. The global changes in gene expression strongly correlated with changes in H3K4me3, but not H3K27me3. High expression of DORMANCY-ASSOCIATED MADS-box (DAM) genes, key regulators of dormancy, in autumn was associated with high H3K4me3 levels. In addition, known DAM/SHORT VEGETATIVE PHASE (SVP) target genes significantly overlapped with H3K4me3-modified genes as bud dormancy progressed. These data suggest that H3K4me3 contributes to the central dormancy circuit, consisting of DAM/SVP and abscisic acid (ABA), in autumn. In winter, the lower expression and H3K4me3 levels at DAMs and gibberellin metabolism genes control chilling-induced release of dormancy. Warming conditions in spring facilitate the expression of genes related to phytohormones, the cell cycle, and cell wall modification by increasing H3K4me3 toward bud break. Our study also revealed that activation of auxin and repression of ABA sensitivity in spring are conditioned at least partly through temperature-mediated epigenetic regulation in winter., 果樹が季節に応答して休眠から目覚めるしくみを解明 --果樹の休眠と発芽におけるヒストン修飾の寄与を解析--. 京都大学プレスリリース. 2022-07-01.

In degraded karst ecosystem, vegetation restoration efforts almost exclusively rely on planted seedlings, but this is not effective to maintain community diversity and resilience. As seed functional traits, seed dormancy and germination are key to community assembly. Unfortunately, these elements are commonly overlooked in restoring degraded ecosystems. This work classifies seed dormancy of 27 species with different life forms that are common on a degraded karst mountain. We examined the effects of temperature regime and light conditions on percentage germination and assessed the relationships between seed traits and germination index using a partial least squares regression (PLSR). Approximately 48% of the investigated species had physiological dormancy, 37% were non-dormant, 7% had morphophysiological dormancy, 4% had morphological dormancy and 4% had physical dormancy. We found that 94% (15 out of 16) species had maximum germination in warm temperature regimes (20/13 and 25/18 °C), while the remaining species required cool temperatures (10/4 °C). PLSR analysis indicated a significant positive correlation between seed mass and T

Background For species that produce seeds with a water-impermeable coat, i.e. physical dormancy (PY), it has been widely recognized that (1) seeds shed at a permeable state cannot become impermeable after dispersal; and (2) dormancy-cycling, i.e. a permeable ↔ impermeable transition, does not occur. Given a tight relationship between moisture content and onset of seed-coat impermeability, seeds maturing at low relative humidity (RH) and occurring in a high-temperature environment are inferred to produce impermeable coats, and ex situ drying of permeable seeds can lead to the onset of impermeability. Scope and conclusion It is proposed here that permeable seeds dispersed at low RH and in high-temperature soils might become impermeable due to continuous drying. Similarly, seeds with shallow PY dormancy (with higher moisture content immediately after becoming impermeable) can cycle back to a permeable state or absolute PY (complete dry state) when RH increases or decreases, respectively. A conceptual model is developed to propose that seeds from several genera of 19 angiosperm families at the time of natural dispersal can be (1) impermeable (dormant), i.e. primary dormancy; (2) impermeable (dormant) and become permeable (non-dormant) and then enter a dormant state in the soil, often referred to as secondary dormancy; (3) permeable (non-dormant) and become impermeable (dormant) in the soil, i.e. enforced dormancy; or (4) dormant or non-dormant, but cycle between permeable and non-permeable states depending on the soil conditions, i.e. dormancy-cycling, which is different from sensitivity-cycling occurring during dormancy break. It is suggested that this phenomenon could influence the dormancy-breaking pattern, but detailed studies of this are lacking.

Plants growing in temperate and boreal regions of the world have to face strikingly different environmental conditions during summer and winter. Being sessile organisms, plants have had to develop various strategies to adapt to these changes in light, temperature, and water availability, thereby optimizing their 'economy of growth'. While annual plants can endure unfavorable winter conditions in the form of a seed, or under a protective cover of thick snow, perennial plants such as trees adapt by going into a stage of deep sleep called winter dormancy. To enter dormancy, vegetative growth is stopped in the late summer or early autumn and the shoots are converted into buds, where the shoot apical meristems are protected by tightly closed and hardened bud scales (Figures 1 and 2). At the same time, cold hardiness develops and the need for water and nutrient uptake is drastically reduced. Deciduous trees also go through leaf senescence whereby the leaves develop their autumn colors and are shed (Figure 1A). The trees then spend the beginning of the winter in a state of deep sleep in which they are completely unreceptive to any environmental signals telling them to wake up. However, as winter progresses, the trees are gradually released from this slumber and will eventually flush their buds in the spring. Vegetative growth then resumes with the formation of new leaves and shoots during summer until the trees again go into growth cessation and the cycle is closed (Figures 1 and 2). This cycle of growth and dormancy is central for the ability of trees to adapt to growth at different latitudes and elevations. The further north, or the higher the elevation at which the trees grow, the earlier in the season the trees enter growth cessation and the later they flush their buds in the spring. This is because meteorological winter arrives earlier in the season and lasts longer into the spring. The trees therefore have to stop growth earlier in the season to ensure that they have enough time to complete bud formation and to develop cold hardiness and dormancy. They also have to be sure that winter is really over before flushing their buds. Winter dormancy is therefore a clear case of a trade-off between the length of the growing season and the protection against winter damage - a nice example of 'economy in biology', the theme of this special issue. This primer will briefly summarize what we know about the environmental signals that influence the annual growth cycle in trees, as well as our current understanding of the genetic pathways and molecular mechanisms regulated by these signals.

Developing innovative agri-technologies is essential for the sustainable intensification of global food production. Seed dormancy is an adaptive trait which defines the environmental conditions in which the seed is able to germinate. Dormancy release requires sensing and integration of multiple environmental signals, a complex process which may be mimicked by seed treatment technologies. Here, we reveal molecular mechanisms by which non-thermal (cold) atmospheric gas plasma-activated water (GPAW) releases the physiological seed dormancy of Arabidopsis thaliana. GPAW triggered dormancy release by synergistic interaction between plasma-generated reactive chemical species (NO3–, H2O2, ·NO, and ·OH) and multiple signalling pathways targeting gibberellin and abscisic acid (ABA) metabolism and the expression of downstream cell wall-remodelling genes. Direct chemical action of GPAW on cell walls resulted in premature biomechanical endosperm weakening. The germination responses of dormancy signalling (nlp8, prt6, and dog1) and ABA metabolism (cyp707a2) mutants varied with GPAW composition. GPAW removes seed dormancy blocks by triggering multiple molecular signalling pathways combined with direct chemical tissue weakening to permit seed germination. Gas plasma technologies therefore improve seed quality by mimicking permissive environments in which sensing and integration of multiple signals lead to dormancy release and germination.

Seed dormancy is an important life history state that increases survival and fitness of seed plants, and thus it has attracted much attention. However, global biogeography, effects of paleoenvironment, evolutionary roles of dormancy transitions, and differences in adaptations of seed dormancy between life-forms are poorly understood. We compiled global distribution records for seed dormancy of 12 743 species and their phylogeny to explore the biogeographic patterns, environmental drivers, and evolutionary transitions between seed dormancy and nondormancy. Biogeographic patterns reveal a low proportion of dormancy in tropical rainforest regions and arctic regions and a high proportion of dormancy in remaining tropical, subtropical, and temperate regions for all species and woody species. Herbaceous plants show a greater proportion of dormancy in most global regions except arctic regions. Seasonal environments have a consistent positive influence on the dormancy pattern for both life-forms, but precipitation and temperature were important driving factors for woody and herbaceous plants, respectively. Seed dormancy was the dominating state during the evolutionary history of seed plants, and dormancy transitions had a significant relationship with paleotemperatures. Dormancy and nondormancy transitions in response to fluctuating environments during long-term evolutionary history may have played important roles in the diversification of seed plants. Our results add to the current knowledge about seed dormancy from macro-adaptive perspectives and the potential adaptive mechanisms of seed plants.

Background and Aims Interactions between ecological factors and seed physiological responses during the establishment phase shape the distribution of plants. Yet, our understanding of the functions and evolution of early-life traits has been limited by the scarcity of large-scale datasets. Here, we tested the hypothesis that the germination niche of temperate plants is shaped by their climatic requirements and phylogenetic relatedness, using germination data sourced from a comprehensive seed conservation database of the European flora (ENSCOBASE). Methods We performed a phylogenetically informed Bayesian meta-analysis of primary data, considering 18 762 germination tests of 2418 species from laboratory experiments conducted across all European geographical regions. We tested for the interaction between species’ climatic requirements and germination responses to experimental conditions including temperature, alternating temperature, light and dormancy-breaking treatments, while accounting for between-study variation related to seed sources and seed lot physiological status. Key Results Climate was a strong predictor of germination responses. In warm and seasonally dry climates the seed germination niche includes a cold-cued germination response and an inhibition determined by alternating temperature regimes and cold stratification, while in climates with high temperature seasonality opposite responses can be observed. Germination responses to scarification and light were related to seed mass but not to climate. We also found a significant phylogenetic signal in the response of seeds to experimental conditions, providing evidence that the germination niche is phylogenetically constrained. Nevertheless, phylogenetically distant lineages exhibited common germination responses under similar climates. Conclusion This is the first quantitative meta-analysis of the germination niche at a continental scale. Our findings showed that the germination niches of European plants exhibit evolutionary convergence mediated by strong pressures at the macroclimatic level. In addition, our methodological approach highlighted how large datasets generated by conservation seed banking can be valuable sources to address questions in plant macroecology and evolution.

PREMISE In temperate sand dunes, rising air temperature owing to climate change could not only further elevate surface soil temperatures during summers but also drastically change the range of soil temperatures in other seasons. Winter warming may shift the timing of seed germination of dune species that require cold stratification for dormancy release. METHODS We assessed the effects of temperature on dormancy and germination of Viola grayi seeds and evaluated whether winter warming could affect its germination phenology by conducting germination experiments and analyzing soil temperature data in cold and warm winters. RESULTS Viola grayi seeds were dormant when dispersed in spring. One-month moist-chilling treatment (4°C) effectively released dormancy, while short, intermittent lower temperatures (alternating 20/5°C) did not. Continuous higher temperatures induced secondary dormancy in non-dormant seeds. During a cold, snowy winter, the surface soil temperatures of the sand dune remained at 0-2°C for approximately one month owing to the accumulated snow, while the period of such stable low soil temperatures was much shorter during a warm, less snowy winter, and the highest soil temperature class reached 20-25°C. These results suggest that dispersed seeds germinate in the following spring after winter chilling, but they may remain dormant after warm winters. CONCLUSIONS With winter warming, prolonged seed dormancy and associated germination delay could occur in V. grayi seeds. Assessing the minimum requirements for dormancy release and the potential to form persistent soil seed banks is important for judging the necessity and urgency of conservation efforts for temperate dune species. This article is protected by copyright. All rights reserved.

The dynamic behaviour of seeds in soil seed banks depends on their ability to act as sophisticated environmental sensors to adjust their sensitivity thresholds for germination by dormancy mechanisms. Here we show that prolonged incubation of sugar beet fruits at low temperature (chilling at 5°C, generally known to release seed dormancy of many species) can induce secondary nondeep physiological dormancy of an apparently nondormant crop species. The physiological and biophysical mechanisms underpinning this cold-induced secondary dormancy include the chilling-induced accumulation of abscisic acid in the seeds, a reduction in the embryo growth potential and a block in weakening of the endosperm covering the embryonic root. Transcriptome analysis revealed distinct gene expression patterns in the different temperature regimes and upon secondary dormancy induction and maintenance. The chilling caused reduced expression of cell wall remodelling protein genes required for embryo cell elongation growth and endosperm weakening, as well as increased expression of seed maturation genes, such as for late embryogenesis abundant proteins. A model integrating the hormonal signalling and master regulator expression with the temperature-control of seed dormancy and maturation programmes is proposed. The revealed mechanisms of the cold-induced secondary dormancy are important for climate-smart agriculture and food security.

Graphical abstract, Highlights • Functional ABA biosynthesis genes show specific roles for ABA accumulation at different stages of seed development and seedling establishment. • De novo ABA biosynthesis during embryogenesis is required for late seed development, maturation, and induction of primary dormancy. • ABA plays multiple roles with the key LAFL hub to regulate various downstream signaling genes in seed and seedling development. • Key ABA signaling genes ABI3, ABI4, and ABI5 play important multiple functions with various cofactors during seed development such as de-greening, desiccation tolerance, maturation, dormancy, and seed vigor. • The crosstalk between ABA and other phytohormones are complicated and important for seed development and seedling establishment., Background Seed is vital for plant survival and dispersion, however, its development and germination are influenced by various internal and external factors. Abscisic acid (ABA) is one of the most important phytohormones that influence seed development and germination. Until now, impressive progresses in ABA metabolism and signaling pathways during seed development and germination have been achieved. At the molecular level, ABA biosynthesis, degradation, and signaling genes were identified to play important roles in seed development and germination. Additionally, the crosstalk between ABA and other hormones such as gibberellins (GA), ethylene (ET), Brassinolide (BR), and auxin also play critical roles. Although these studies explored some actions and mechanisms by which ABA-related factors regulate seed morphogenesis, dormancy, and germination, the complete network of ABA in seed traits is still unclear. Aim of review Presently, seed faces challenges in survival and viability. Due to the vital positive roles in dormancy induction and maintenance, as well as a vibrant negative role in the seed germination of ABA, there is a need to understand the mechanisms of various ABA regulators that are involved in seed dormancy and germination with the updated knowledge and draw a better network for the underlying mechanisms of the ABA, which would advance the understanding and artificial modification of the seed vigor and longevity regulation. Key scientific concept of review Here, we review functions and mechanisms of ABA in different seed development stages and seed germination, discuss the current progresses especially on the crosstalk between ABA and other hormones and signaling molecules, address novel points and key challenges (e.g., exploring more regulators, more cofactors involved in the crosstalk between ABA and other phytohormones, and visualization of active ABA in the plant), and outline future perspectives for ABA regulating seed associated traits.

During the course of evolution, different ecotypes of rice (Oryza sativa L.) have evolved distinct strategies to cope with submergence stress. Such contrasting responses are mediated by plant hormones that are principle regulators of growth, development and responses to various biotic and abiotic stresses. These hormones act cooperatively and show extensive crosstalk which is mediated by key regulatory genes that serve as nodes of molecular communication. The presence or absence of such genes leads to significant changes in hormone signalling pathways and hence, governs the type of response that the plant will exhibit. As flooding is one of the leading causes of crop loss across all the major rice-producing countries, it is crucial to deeply understand the molecular nexus governing the response to submergence to produce flood resilient varieties. This review focuses on the hormonal signalling pathways that mediate two contrasting responses of the rice plant to submergence stress namely, rapid internode elongation to escape flood waters and quiescence response that enables the plant to survive under complete submergence. The significance of several key genes such as Sub1A-1, SLR1, SD1 and SK1/SK2, in defining the ultimate response to submergence has also been discussed.

GATA transcription factors have been reported to function in plant growth and development and during various biotic/abiotic stresses in Arabidopsis and rice. However, the functions of wheat GATAs, particularly in the regulation of seed dormancy and germination, remain unclear. Here, we identified 78 TaGATAs in wheat and divided them into five subfamilies. Sixty-four paralogous pairs and 52 orthologous pairs were obtained, and Ka/Ks ratios showed that the TaGATAs had undergone strong purifying election during the evolutionary process. Triplet analysis indicated that a high homologue retention rate could explain the large number of TaGATAs in wheat. Gene structure analysis revealed that most members of the same subfamily had similar structures, and subcellular localization prediction indicated that most TaGATAs were located in the nucleus. Gene ontology annotation results showed that most TaGATAs had molecular functions in DNA and zinc binding, and promoter analysis suggested that they may play important roles in growth, development, and biotic/abiotic stress response. We combined three microarray datasets with qRT-PCR expression data from wheat varieties of contrasting dormancy and pre-harvest sprouting resistance levels during imbibition in order to identify ten candidate genes (TaGATA17/-25/-34/-37/-40/-46/-48/-51/-72/-73) that may be involved in the regulation of seed dormancy and germination in wheat. These findings provide valuable information for further dissection of TaGATA functions in the regulation of seed dormancy and germination, thereby enabling the improvement of wheat pre-harvest sprouting resistance by gene pyramiding.

Underdeveloped (small) embryos embedded in abundant endosperm tissue, and thus having morphological dormancy (MD) or morphophysiological dormancy (MPD), are considered to be the ancestral state in seed dormancy evolution. This trait is retained in the Apiaceae family, which provides excellent model systems for investigating the underpinning mechanisms. We investigated Apium graveolens (celery) MD by combined innovative imaging and embryo growth assays with the quantification of hormone metabolism, as well as the analysis of hormone and cell-wall related gene expression. The integrated experimental results demonstrated that embryo growth occurred inside imbibed celery fruits in association with endosperm degradation, and that a critical embryo size was required for radicle emergence. The regulation of these processes depends on gene expression leading to gibberellin and indole-3- acetic acid (IAA) production by the embryo and on crosstalk between the fruit compartments. ABA degradation associated with distinct spatiotemporal patterns in ABA sensitivity control embryo growth, endosperm breakdown and radicle emergence. This complex interaction between gibberellins, IAA and ABA metabolism, and changes in the tissuespecific sensitivities to these hormones is distinct from non-MD seeds. We conclude that the embryo growth to reach the critical size and the associated endosperm breakdown inside MD fruits constitute a unique germination programme.

Available studies are far from giving a complete figure for the germination requirements of many Mediterranean Basin species. In this study, we investigated the germination properties of Lavandula stoechas L. (Lamiaceae) in response to different light and temperature regimes. We performed comprehensive experiments to test the effect of fixed and alternating temperatures, dark versus light conditions, and thermodormancy or thermoinhibition on several germination properties in three populations of L. stoechas from southwestern Turkey. Germination patterns showed a substantial variation among populations. Germination percentage was higher in the light conditions than in the dark at most temperatures, and alternating temperatures substituted light for the germination. The requirement of L. stoechas seeds to light for germination gradually increased through lower and higher temperatures than 15 °C. High temperature (30 °C) reduced the germination percentage to zero. However, in the presence of light, seeds were able to germinate after transferring to the optimum temperature (thermoinhibition), but this was not the case in darkness (thermodormancy). Seed germination in L. stoechas was stimulated by light, suggesting that germination has a phytochrome-mediated response. Our study makes an inference to the fire-environment interactions in the germination of Mediterranean seeder species by showing that both light and alternating temperatures stimulate germination in L. stoechas, a species well-known with its fire-related germination. Comprehensive germination studies are required for a better understanding of the early life-stage adaptations of plants to Mediterranean conditions, and for developing more robust strategies for conservation and ecosystem restoration.

Summary High humidity during harvest season often causes pre‐harvest sprouting in barley (Hordeum vulgare). Prolonged grain dormancy prevents pre‐harvest sprouting; however, extended dormancy can interfere with malt production and uniform germination upon sowing. In this study, we used Cas9‐induced targeted mutagenesis to create single and double mutants in QTL FOR SEED DORMANCY 1 (Qsd1) and Qsd2 in the same genetic background. We performed germination assays in independent qsd1 and qsd2 single mutants, as well as in two double mutants, which revealed a strong repression of germination in the mutants. These results demonstrated that normal early grain germination requires both Qsd1 and Qsd2 function. However, germination of qsd1 was promoted by treatment with 3% hydrogen peroxide, supporting the notion that the mutants exhibit delayed germination. Likewise, exposure to cold temperatures largely alleviated the block of germination in the single and double mutants. Notably, qsd1 mutants partially suppress the long dormancy phenotype of qsd2, while qsd2 mutant grains failed to germinate in the light, but not in the dark. Consistent with the delay in germination, abscisic acid accumulated in all mutants relative to the wild type, but abscisic acid levels cannot maintain long‐term dormancy and only delay germination. Elucidation of mutant allele interactions, such as those shown in this study, are important for fine‐tuning traits that will lead to the design of grain dormancy through combinations of mutant alleles. Thus, these mutants will provide the necessary germplasm to study grain dormancy and germination in barley.

Light is considered a factor that influences the seed germination of many weed species, and it can signal whether the environmental conditions are favorable or are not favorable for germination. We aimed to study if there is an influence of light quality and dormancy overcoming in seed germination of Echium plantagineum L. We carried out a 2 x 6 factorial experiment, with and without dormancy overcoming with potassium nitrate followed by immersion in gibberellic acid; six light qualities, obtained through the light filters: blue, green, red, far-red, white light and absence of light. The evaluations performed were germination speed index (GSI), average germination time (AGT), germination at the four and 14 days after seeding (DAS), accumulated germination and relative frequency of germination. We observed significant interaction among the light qualities and seed dormancy overcoming or not for the studied variables. There was no significant effect of light qualities, in the evaluated variables, when performing dormancy overcoming, presenting germination above 90% in all the light qualities. However, without dormancy overcoming, we observed greater GSI, germination at four and 14 DAS for the red light filter with 5, 4, 29 and 45%, respectively. When the seeds were submitted to the absence of light, and without dormancy overcoming, there was only 7% of germination at 14 DAS. The seeds of E. plantagineum presented greater germination under incidence of red light, without dormancy overcoming, being classified as preferably positively photoblastics, provided that the dormancy is not overcome. Resumo A luz é considerada um fator que influencia a germinação das sementes de muitas espécies de plantas daninhas, podendo sinalizar se as condições ambientais são favoráveis ou não para a germinação. Objetivou-se estudar se há influência da qualidade da luz e superação de dormência na germinação de sementes de Echium plantagineum L. Realizou-se um experimento fatorial 2 x 6, com e sem superação de dormência com nitrato de potássio seguido pela imersão em ácido giberélico; seis qualidades de luz, obtidas através de filtros de luz: azul, verde, vermelho, vermelho-distante, luz branca e ausência de luz. As avaliações realizadas foram índice de velocidade de germinação (IVG), tempo médio de germinação (TMG), germinação aos quatro e 14 dias após a semeadura (DAS), germinação acumulada e frequência relativa de germinação. Observou-se interação significativa entre as qualidades de luz e a superação ou não de dormência das sementes para as variáveis estudadas. Não houve efeito significativo das qualidades de luz, nas variáveis avaliadas, ao realizar superação de dormência, apresentando germinação acima de 90% em todas as qualidades de luz. Todavia, sem superação de dormência, observou-se maior IVG, germinação aos quatro e 14 DAS para o filtro de luz vermelha com 5,4, 29 e 45%, respectivamente. Quando as sementes foram submetidas à ausência de luz, e sem superação de dormência, houve apenas 7% de germinação aos 14 DAS. As sementes de E. plantagineum apresentam maior germinação sob incidência de luz vermelha, sem superação de dormência, sendo classificadas como fotoblásticas positivas preferenciais, desde que não seja superada a dormência.

Alternative splicing (AS) plays a crucial role in plant growth, development and response to various environmental changes. However, whether alternative splicing of MADS-box transcription factors contributes to the flower bud dormancy process in fruit trees still remains unknown. In this work, the AS profile of genes in the dormant flower buds of 'Dangshansu' pear tree were examined. A total number of 3661 alternatively spliced genes were identified, and three mRNA isoforms of the dormancy associated MADS box (DAM) gene, PpDAM1, derived by alternative splicing, designated as PpDAM1.1, PpDAM1.2 and PpDAM1.3, were characterized. Bimolecular fluorescence complementation (BiFC) analysis indicated that AS of PpDAM1 didn't affect the nucleus localization and homo-/heterodimerization of PpDAM1.1, PpDAM1.2 and PpDAM1.3 proteins, but disturbed the translocation of PpDAM1.1/PpDAM1.1, PpDAM1.3/PpDAM1.3, PpDAM1.1/PpDAM1.3, and PpDAM1.2/PpDAM1.3 dimers to the nucleus. Constitutive expression of PpDAM1.2, but not PpDAM1.1 and PpDAM1.3, in Arabidopsis retarded the growth and development of transgenic plants. Further comparative expression analyses of PpDAM1.1, PpDAM1.2 and PpDAM1.3 in the flower buds of 'Dangshansu' and a less dormant pear cultivar, 'Cuiguan', exhibited that the expression of all the three isoforms in 'Dangshansu' were significantly higher than in 'Cuiguan', especially PpDAM1.2, which showed a predominantly higher expression than PpDAM1.1 and PpDAM1.3 in both cultivars. Our results suggest that alternative splicing of PpDAM1 could play a crucial role in pear flower bud dormancy process.

Temperature variation during seed set is an important modulator of seed dormancy and impacts the performance of crop seeds through effects on establishment rate. It remains unclear how changing temperature during maturation leads to dormancy and growth vigour differences in nondormant seedlings. Here we take advantage of the large seed size in Brassica oleracea to analyse effects of temperature on individual seed tissues. We show that warm temperature during seed maturation promotes seed germination, while removal of the endosperm from imbibed seeds abolishes temperature-driven effects on germination. We demonstrate that cool temperatures during early seed maturation lead to abscisic acid (ABA) retention specifically in the endosperm at desiccation. During this time temperature affects ABA dynamics in individual seed tissues and regulates ABA catabolism. We also show that warm-matured seeds preinduce a subset of germination-related programmes in the endosperm, whereas cold-matured seeds continue to store maturation-associated transcripts including DOG1 because of effects on mRNA degradation before quiescence, rather than because of the effect of temperature on transcription. We propose that effects of temperature on seed vigour are explained by endospermic ABA breakdown and the divergent relationships between temperature and mRNA breakdown and between temperature, seed moisture and the glass transition.

Background To adapt seasonal climate changes under natural environments, Polygonatum sibiricum seeds have a long period of epicotyl morphophysiological dormancy, which limits their wide-utilization in the large-scale plant progeny propagation. It has been proven that the controlled consecutive warm and cold temperature treatments can effectively break and shorten this seed dormancy status to promote its successful underdeveloped embryo growth, radicle emergence and shoot emergence. To uncover the molecular basis of seed dormancy release and seedling establishment, a SMRT full-length sequencing analysis and an Illumina sequencing-based comparison of P. sibiricum seed transcriptomes were combined to investigate transcriptional changes during warm and cold stratifications. Results A total of 87,251 unigenes, including 46,255 complete sequences, were obtained and 77,148 unigenes (88.42%) were annotated. Gene expression analyses at four stratification stages identified a total of 27,059 DEGs in six pairwise comparisons and revealed that more differentially expressed genes were altered at the Corm stage than at the other stages, especially Str_S and Eme. The expression of 475 hormone metabolism genes and 510 hormone signaling genes was modulated during P. sibiricum seed dormancy release and seedling emergence. One thousand eighteen transcription factors and five hundred nineteen transcription regulators were detected differentially expressed during stratification and germination especially at Corm and Str_S stages. Of 1246 seed dormancy/germination known DEGs, 378, 790, and 199 DEGs were associated with P. sibiricum MD release (Corm vs Seed), epicotyl dormancy release (Str_S vs Corm), and the seedling establishment after the MPD release (Eme vs Str_S). Conclusions A comparison with dormancy- and germination-related genes in Arabidopsis thaliana seeds revealed that genes related to multiple plant hormones, chromatin modifiers and remodelers, DNA methylation, mRNA degradation, endosperm weakening, and cell wall structures coordinately mediate P. sibiricum seed germination, epicotyl dormancy release, and seedling establishment. These results provided the first insights into molecular regulation of P. sibiricum seed epicotyl morphophysiological dormancy release and seedling emergence. They may form the foundation of future studies regarding gene interaction and the specific roles of individual tissues (endosperm, newly-formed corm) in P. sibiricum bulk seed dormancy.

Plant hybridization is a pathway for the evolution of adaptive traits. However, hybridization between adapted and nonadapted populations may affect the persistence of combinations of adaptive alleles evolved through natural selection. Seed dormancy is an adaptive trait for weedy rice because it regulates the timing of seed germination and the persistence of the soil seed bank. Hybridization between weedy and cultivated rice has been confirmed with an adaptive introgression of deep seed dormancy alleles from cultivated rice. Here, we explored the influence of hybridization on the conservation of adaptive allele combinations by evaluating natural variation and genetic structure in seed dormancy-associated genomic regions. Based on sequence variation in the genomic regions associated with seed dormancy, hybrid-derived weedy rice strains maintained most of the adaptive combinations for this trait observed in the parental weedy rice, despite equal representation of the parental weedy and cultivated rice in the whole genome sequence. Moreover, hybrid-derived weedy rice strains were more dormant than their parental weedy rice strains, and this trait was strongly influenced by the environment. This study suggests that hybridization between weedy rice (adaptive allelic combinations for seed dormancy) and cultivated rice (nonadaptive combinations) generates weedy rice strains expressing deep seed dormancy caused by genome stabilization through the removal of alleles derived from cultivated rice, in addition to the adaptive introgression of deep seed dormancy alleles derived from cultivated rice. Thus, hybridization between adapted and nonadapted populations appears to be reinforcing the trajectory towards the evolution of adaptive traits.

Winter dormancy is a protective survival strategy for plants to resist harsh natural environments. In the context of global warming, the progression of dormancy has been significantly affected in perennials, which requires further research. Here, a systematic study was performed to compare the induction of dormancy in two closely related iris species with an ecodormancy-only process, the evergreen Iris japonica Thunb. and the deciduous Iris tectorum Maxim. under artificial conditions. Firstly, morphological and physiological observations were evaluated to ensure the developmental status of the two iris species. Furthermore, the expression patterns of the genes involved in key pathways related to plant winter dormancy were determined, and correlation analyses with dormancy marker genes were conducted. We found that deciduous iris entered dormancy earlier than evergreen iris under artificial dormancy induction conditions. Phytohormones and carbohydrates play roles in coordinating growth and stress responses during dormancy induction in both iris species. Moreover, dormancy-related MADS-box genes and SnRKs (Snf1-related protein kinase) might represent a bridge between carbohydrate and phytohormone interaction during iris dormancy. These findings provide a hypothetical model explaining the later dormancy in evergreen iris compared with deciduous iris under artificial dormancy induction conditions and reveal some candidate genes. The findings of this study could provide new insights into the research of dormancy in perennial plants with an ecodormancy-only process and contribute to effectively managing iris production, postharvest storage, and shipping.

Seed dormancy is an adaptive trait that enables plants to survive adverse conditions and restart growth in a season and location suitable for vegetative and reproductive growth. Control of seed dormancy is also important for crop production and food quality because it can help induce uniform germination and prevent preharvest sprouting. Rice preharvest sprouting quantitative trait locus analysis has identified Seed dormancy 4 (Sdr4) as a positive regulator of dormancy development. Here, we analyzed the loss-of-function mutant of the Arabidopsis ortholog, Sdr4 Like1 (SFL1), and found that the sfl1-1 seeds showed precocious germination at the mid- to late-maturation stage similar to rice sdr4 mutant, but converted to become more dormant than the wild type during maturation drying. Coordinated with the dormancy levels, expression levels of the seed maturation and dormancy master regulator genes, ABI3, FUS3, and DOG1 in sfl1-1 seeds were lower than in wild type at early- and mid-maturation stages, but higher at the late-maturation stage. In addition to the seed dormancy phenotype, sfl1-1 seedlings showed a growth arrest phenotype and heterochronic expression of LAFL (LEC1, ABI3, FUS3, LEC2) and DOG1 in the seedlings. These data suggest that SFL1 is a positive regulator of initiation and termination of the seed dormancy program. We also found genetic interaction between SFL1 and the SFL2, SFL3, and SFL4 paralogs of SFL1, which impacts on the timing of the phase transition from embryo maturation to seedling growth.