Evolutionary origin and functional investigation of the widely conserved plant PEBP gene STEPMOTHER OF FT AND TFL1 (SMFT).

Genes of the family PHOSPHATIDYLETHANOLAMINE-BINDING PROTEINS (PEBP) have been intensely studied in plants for their role in cell (re)programming and meristem differentiation. Recently, sporadic reports of the presence of a new type of PEBP in plants became available, highly similar to the YY-PEBPs of prokaryotes. A comprehensive investigation of their spread, origin, and function revealed conservation across the plant kingdom. The YY-PEBP clade in plants seems to have resulted from a single Horizontal Gene Transfer (HGT) episode from a prokaryotic organism to an ancestral streptophyte. YY-PEBPs are also present in other eukaryotes, such as certain fungi, diatoms, and rotifers, and these cases derive from independent HGT events. Reciprocally, the occurrence of the eukaryotic CETS/RKIP type PEBPs (CR-PEBPs) was noticed in bacteria of the genus Nocardia, showing that HGT has occurred as well from eukaryotes to prokaryotes. Based on these observations, we propose that the current model of the PEBP family in plants needs to be updated with the clade STEPMOTHER OF FT AND TFL1 (SMFT). SMFT genes not only share high sequence conservation but also show specific expression in homologous plant structures that serve as propagules. Functional analysis of Arabidopsis smft mutant lines pointed to a function for this gene in regulating seed germination, both concerning primary dormancy release and in response to adverse high-temperature conditions. Overall, our study reveals an increasing complexity in the evolutionary history of the PEBP gene family, unlocking new potential in understanding the evolution and functional spectrum of these important key regulatory genes.

Current Insights into Weak Seed Dormancy and Pre-Harvest Sprouting in Crop Species.

During the domestication of crops, seed dormancy has been reduced or eliminated to encourage faster and more consistent germination. This alteration makes cultivated crops particularly vulnerable to pre-harvest sprouting, which occurs when mature crops are subjected to adverse environmental conditions, such as excessive rainfall or high humidity. Consequently, some seeds may bypass the normal dormancy period and begin to germinate while still attached to the mother plant before harvest. Grains affected by pre-harvest sprouting are characterized by increased levels of α-amylase activity, resulting in poor processing quality and immediate grain downgrading. In the agriculture industry, pre-harvest sprouting causes annual economic losses exceeding USD 1 billion worldwide. This premature germination is influenced by a complex interplay of genetic, biochemical, and molecular factors closely linked to environmental conditions like rainfall. However, the exact mechanism behind this process is still unclear. Unlike pre-harvest sprouting, vivipary refers to the germination process and the activation of α-amylase during the soft dough stage, when the grains are still immature. Mature seeds with reduced levels of ABA or impaired ABA signaling (weak dormancy) are more susceptible to pre-harvest sprouting. While high seed dormancy can enhance resistance to pre-harvest sprouting, it can lead to undesirable outcomes for most crops, such as non-uniform seedling establishment after sowing. Thus, resistance to pre-harvest sprouting is crucial to ensuring productivity and sustainability and is an agronomically important trait affecting yield and grain quality. On the other hand, seed color is linked to sprouting resistance; however, the genetic relationship between both characteristics remains unresolved. The identification of mitogen-activated protein kinase kinase-3 (MKK3) as the gene responsible for pre-harvest sprouting-1 (Phs-1) represents a significant advancement in our understanding of how sprouting in wheat is controlled at the molecular and genetic levels. In seed maturation, Viviparous-1 (Vp-1) plays a crucial role in managing pre-harvest sprouting by regulating seed maturation and inhibiting germination through the suppression of α-amylase and proteases. Vp-1 is a key player in ABA signaling and is essential for the activation of the seed maturation program. Mutants of Vp-1 exhibit an unpigmented aleurone cell layer and exhibit precocious germination due to decreased sensitivity to ABA. Recent research has also revealed that TaSRO-1 interacts with TaVp-1, contributing to the regulation of seed dormancy and resistance to pre-harvest sprouting in wheat. The goal of this review is to emphasize the latest research on pre-harvest sprouting in crops and to suggest possible directions for future studies.

Soil Seed Bank of the Alpine Endemic Carnation, Dianthus pavonius Tausch (Piedmont, Italy), a Useful Model for the Study of Host-Pathogen Dynamics.

Soil seedbanks are particularly important for the resiliency of species living in habitats threatened by climate change, such as alpine meadows. We investigated the germination rate and seedbank potential for the endemic species Dianthus pavonius, a carnation native to the Maritime Alps that is used as model system for disease in natural populations due to its frequent infections by a sterilizing anther-smut pathogen. We aimed to ascertain whether this species can create a persistent reserve of viable seeds in the soil which could impact coevolutionary dynamics. Over three years, we collected data from seeds sown in natural soil and analyzed their germination and viability. We found that D. pavonius seeds are not physiologically dormant and they are able to create a persistent soil seed bank that can store seeds in the soil for up to three years, but lower than the estimated plant lifespan. We conclude that while the seedbank may provide some demographic stability to the host population, its short duration is unlikely to strongly affect the host's ability to respond to selection from disease. Our findings have implications for the conservation of this alpine species and for understanding the evolutionary dynamics between the host and its pathogen.

The MKK3 module integrates nitrate and light signals to modulate secondary dormancy in Arabidopsis thaliana.

Seed dormancy corresponds to a reversible blockage of germination. Primary dormancy is established during seed maturation, while secondary dormancy is set up on the dispersed seed, following an exposure to unfavorable factors. Both dormancies are relieved in response to environmental factors, such as light, nitrate, and coldness. Quantitive Trait Locus (QTL) analyses for preharvest sprouting identified MKK3 kinase in cereals as a player in dormancy control. Here, we showed that MKK3 also plays a role in secondary dormancy in Arabidopsis within a signaling module composed of MAP3K13/14/19/20, MKK3, and clade-C MAPKs. Seeds impaired in this module acquired heat-induced secondary dormancy more rapidly than wild-type (WT) seeds, and this dormancy is less sensitive to nitrate, a signal able to release dormancy. We also demonstrated that MPK7 was strongly activated in the seed during dormancy release, especially in response to light and nitrate. This activation was greatly reduced in map3k13/14/19/20 and mkk3 mutants. Finally, we showed that the module was not regulated and apparently did not regulate the genes controlling abscisic acid/gibberellin acid hormone balance, one of the crucial mechanisms of seed dormancy control. Overall, our work identified a MAPK module controlling seed germination and enlarged the panel of functions of the MKK3-related modules in plants.

Population histories of variable reproductive success and low winter precipitation correlate with risk-averse seed germination in a mediterranean-climate winter annual.

PREMISE: Seed germination involves risk; post-germination conditions might not allow survival and reproduction. Variable, stressful environments favor seeds with germination that avoids risk (e.g., germination in conditions predicting success), spreads risk (e.g., dormancy), or escapes risk (e.g., rapid germination). Germination studies often investigate trait correlations with climate features linked to variation in post-germination reproductive success. Rarely are long-term records of population reproductive success available. METHODS: Supported by demographic and climate monitoring, we analyzed germination in the California winter-annual Clarkia xantiana subsp. xantiana. Sowing seeds of 10 populations across controlled levels of water potential and temperature, we estimated temperature-specific base water potential for 20% germination, germination time weighted by water potential above base (hydrotime), and a dormancy index (frequency of viable, ungerminated seeds). Mixed-effects models analyzed responses to (1) temperature, (2) discrete variation in reproductive success (presence or absence of years with zero seed production by a population), and (3) climate covariates, mean winter precipitation and coefficient of variation (CV) of spring precipitation. For six populations, records enabled analysis with a continuous metric of variable reproduction, the CV of per-capita reproductive success. RESULTS: Populations with more variable reproductive success had higher base water potential and dormancy. Higher base water potential and faster germination occurred at warmer experimental temperatures and in seeds of populations with wetter winters. CONCLUSIONS: Geographic variation in seed germination in this species suggests local adaptation to demographic risk and rainfall. High base water potential and dormancy may concentrate germination in years likely to allow reproduction, while spreading risk among years.

Seed functional ecology in Brazilian rock outcrop vegetation: an integrative synthesis.

BACKGROUND AND AIMS: Rock outcrop vegetation is distributed worldwide and hosts a diverse and unique flora that evolved under harsh environmental conditions. Unfortunately, seed ecology in such ecosystems has received little attention, especially regarding seed traits, germination responses to abiotic factors and the potential role of phylogenetic relatedness on such features Here, we provide the first quantitative and phylogenetically-informed synthesis of the seed functional ecology of Brazilian rock outcrop vegetation, with a particular focus on quartzitic and ironstone campo rupestre. METHODS: Using a database of functional trait data, we calculated the phylogenetic signal of seven seed traits for 371 taxa and tested whether they varied among growth forms, geographic distribution, and microhabitats. We also conducted meta-analyses that included 4,252 germination records for 102 taxa to assess the effects of light, temperature, and fire-related cues on the germination of campo rupestre species and explored how the aforementioned ecological groups and seed traits modulate germination responses. KEY RESULTS: All traits and germination responses showed a moderate-to-strong phylogenetic signal. Campo rupestre species responded positively to light and had maximum germination between 20-25 ºC. The effect of temperatures beyond this range was moderated by growth form, species geographic distribution, and microhabitat. Seeds exposed to heat shocks above 80 °C lost viability, but smoke accelerated germination. We found a moderating effect of seed mass for in responses to light and heat shocks, with larger, dormant seeds tolerating heat better but less sensitive to light. Species from xeric habitats evolved phenological strategies to synchronise germination during periods of increased soil water availability. CONCLUSIONS: Phylogenetic relatedness plays a major role in shaping seed ecology of Brazilian rock outcrop vegetation. Nevertheless, seed traits and germination responses varied significantly between growth forms, species geographic distribution and microhabitats, providing support to the regeneration niche hypothesis and the role of functional traits in shaping germination in these ecosystems.

Characterizing seed dormancy in Epimedium brevicornu Maxim.: Development of novel chill models and determination of dormancy release mechanisms by transcriptomics.

PURPOSE: Epimedium brevicornu Maxim. is a perennial persistent C3 plant of the genus Epimedium Linn. in the family Berberaceae that exhibits severe physiological and morphological seed dormancy.We placed mature E. brevicornu seeds under nine stratification treatment conditions and explored the mechanisms of influence by combining seed embryo growth status assessment with related metabolic pathways and gene co-expression analysis. RESULTS: We identified 3.9 °C as the optimum cold-stratification temperature of E. brevicornu seeds via a chilling unit (CU) model. The best treatment was variable-temperature stratification (10/20 °C, 12/12 h) for 4 months followed by low-temperature stratification (4 °C) for 3 months (4-3). A total of 63801 differentially expressed genes were annotated to 2587 transcription factors (TFs) in 17 clusters in nine treatments (0-0, 0-3, 1-3, 2-3, 3-3, 4-3, 4-2, 4-1, 4-0). Genes specifically highly expressed in the dormancy release treatment group were significantly enriched in embryo development ending in seed dormancy and fatty acid degradation, indicating the importance of these two processes. Coexpression analysis implied that the TF GRF had the most reciprocal relationships with genes, and multiple interactions centred on zf-HD and YABBY as well as on MYB, GRF, and TCP were observed. CONCLUSION: In this study, analyses of plant hormone signal pathways and fatty acid degradation pathways revealed changes in key genes during the dormancy release of E. brevicornu seeds, providing evidence for the filtering of E. brevicornu seed dormancy-related genes.

Germination Assays for Comparable Seed Dormancy Depth and Distinction of Seed Color by Multiplex PCR in Wheat.

Seed dormancy is an important trait in cereal breeding, as it prevents preharvest sprouting (PHS). Although seed dormancy is a multifactorial trait, seed color has been demonstrated to be a major dormancy-related factor controlled by few genes. The R-1 gene is a seed color regulator that encodes a MYB-type transcription factor in wheat. A set of genetic markers designed against R-1 can provide a powerful tool for swift wheat breeding. Depth of seed dormancy varies not only among lines but also during seed development in each line. In this chapter, we describe how developmental seeds can be collected to perform germination tests, how seed color can be observed after NaOH staining, and how to genotype wheat R-1 genes using multiplex PCR.

Map-Based Cloning of the Causal Gene for a Seed Dormancy Quantitative Trait Locus in Barley.

Seed dormancy is an important agronomic trait in cereal crops. Throughout the domestication of cereals, seed dormancy has been reduced to obtain uniform germination. However, grain crops must retain moderate levels of seed dormancy to prevent problems such as preharvest sprouting in wheat (Triticum aestivum) and barley (Hordeum vulgare). To produce modern cultivars with the appropriate seed dormancy levels, it is important to identify the genes responsible for seed dormancy. With recent advances in sequencing technology, several causal genes for seed dormancy quantitative trait loci (QTLs) have been identified in barley and wheat. Here, we present a method to identify causal genes for seed dormancy QTLs in barley, a method that is also applicable to other cereals.

Genome-Wide Association Study of Preharvest Sprouting in Wheat.

Genome-wide association study (GWAS) is widely used to characterize genes or quantitative trait loci (QTLs) associated with preharvest sprouting and seed dormancy. GWAS can identify both previously discovered and novel QTLs across diverse genetic panels. The high-throughput SNP arrays or next-generation sequencing technologies have facilitated the identification of numerous genetic markers, thereby significantly enhancing the resolution of GWAS. Although various methods have been developed, the fundamental principles underlying these techniques remain constant. Here, we provide a basic technological flow to perform seed dormancy assay, followed by GWAS using population structure control, and compared it with previous identified QTLs and genes.

Evaluation Methods for Preharvest Sprouting Resistance in Japanese Wheat Breeding Programs.

Different methodologies have been applied for the selection of preharvest sprouting resistance in cereal breeding programs. We describe here a series of methods used in practical wheat breeding programs in Japan, including phenotyping based on germination score after artificial rain treatments and genotyping using DNA markers. These methods can be modified and applied to breeding programs in which preharvest sprouting is a problem during cereal cultivation.

OsABA3 is Crucial for Plant Survival and Resistance to Multiple Stresses in Rice.

Preharvest sprouting (PHS) is a serious problem in rice production as it leads to reductions in grain yield and quality. However, the underlying mechanism of PHS in rice remains unclear. In this study, we identified and characterized a preharvest sprouting and seedling lethal (phssl) mutant. The heterozygous phssl/+ mutant exhibited normal plant development, but severe PHS in paddy fields. However, the homozygous phssl mutant was seedling lethal. Gene cloning and genetic analysis revealed that a point mutation in OsABA3 was responsible for the mutant phenotypes. OsABA3 encodes a molybdenum cofactor (Moco) sulfurase. The activities of the sulfureted Moco-dependent enzymes such as aldehyde oxidase (AO) and xanthine dehydrogenase (XDH) were barely detectable in the phssl mutant. As the final step of abscisic acid (ABA) de novo biosynthesis is catalyzed by AO, it indicated that ABA biosynthesis was interrupted in the phssl mutant. Exogenous application of ABA almost recovered seed dormancy of the phssl mutant. The knock-out (ko) mutants of OsABA3 generated by CRISPR-Cas9 assay, were also seedling lethal, and the heterozygous mutants were similar to the phssl/+ mutant showing reduced seed dormancy and severe PHS in paddy fields. In contrast, the OsABA3 overexpressing (OE) plants displayed a significant increase in seed dormancy and enhanced plant resistance to PHS. The AO and XDH activities were abolished in the ko mutants, whereas they were increased in the OE plants. Notably, the Moco-dependent enzymes including nitrate reductase (NR) and sulfite oxidase (SO) showed reduced activities in the OE plants. Moreover, the OE plants exhibited enhanced resistances to osmotic stress and bacterial blight, and flowered earlier without any reduction in grain yield. Taken together, this study uncovered the crucial functions of OsABA3 in Moco sulfuration, plant development, and stress resistance, and suggested that OsABA3 is a promising target gene for rice breeding.

Germination response to winter temperature changes with seed shape and length of temperature exposure.

In many regions, the climate is changing faster during winter than during the other seasons, and a loss of snow cover combined with increased temperature variability can expose overwintering organisms to harmful conditions. Understanding how species respond to these changes during critical developmental times, such as seed germination, helps us assess the ecological implications of winter climate change. To address this concern, we measured the breaking of seed dormancy and cold tolerance of temperate grassland species in the lab and field. In the lab, we ran germination trials testing the tolerance of 17 species to an extreme cold event. In the field, we deployed seeds of two species within a snow manipulation experiment at three locations and measured germination success biweekly from seeds subjected to ambient and reduced snow cover from winter into spring. From lab trials, cold tolerance varied among species, with seed germination decreasing <10%-100% following extreme cold events. Cold tolerance was related to seed traits, specifically less round seeds, seeds that required cold stratification, and seeds that mature later in the season tended to be more impacted by extreme cold temperatures. This variation in seed cold tolerance may contribute to altered community composition with continued winter climate change. In the field, germination increased through late winter, coinciding with the accumulation of days where temperatures were favorable for cold stratification. Through spring, germination success decreased as warm temperatures accumulated. Collectively, species-specific seed cold tolerances and mortality rates may contribute to compositional changes in grasslands under continued winter climate change.

Molecular mechanism of vivipary as revealed by the genomes of viviparous mangroves and non-viviparous relatives.

Vivipary is a prominent feature of mangroves, allowing seeds to complete germination while attached to the mother plant, and equips propagules to endure and flourish in challenging coastal intertidal wetlands. However, vivipary-associated genetic mechanisms remain largely elusive. Genomes of two viviparous mangrove species and a non-viviparous inland relative were sequenced and assembled at the chromosome level. Comparative genomic analyses between viviparous and non-viviparous genomes revealed that DELAY OF GERMINATION 1 (DOG1) family genes (DFGs), the proteins from which are crucial for seed dormancy, germination, and reserve accumulation, are either lost or dysfunctional in the entire lineage of true viviparous mangroves but are present and functional in their inland, non-viviparous relatives. Transcriptome dynamics at key stages of vivipary further highlighted the roles of phytohormonal homeostasis, proteins stored in mature seeds, and proanthocyanidins in vivipary under conditions lacking DFGs. Population genomic analyses elucidate dynamics of syntenic regions surrounding the missing DFGs. Our findings demonstrated the genetic foundation of constitutive vivipary in Rhizophoraceae mangroves.

SlWRKY37 targets SlLEA2 and SlABI5-like7 to regulate seed germination vigor in tomato.

Seed germination is a critical phase for the life cycle and propagation of higher plants. This study explores the role of SlWRKY37, a WRKY transcription factor in tomato, in modulating seed germination. We discovered that SlWRKY37 expression is markedly downregulated during tomato seed germination. Through CRISPR/Cas9-mediated editing, we demonstrate that SlWRKY37 knockout enhances germination, while its overexpression results in a delay compared to the wild type. Transcriptome analysis revealed 679 up-regulated and 627 down-regulated genes in Slwrky37-CRISPR deletion mutants relative to the wild type. Gene ontology (GO) enrichment analysis indicated these differentially expressed genes are linked to seed dormancy, abscisic acid homeostasis, and protein phosphorylation pathways. Bioinformatics and biochemical assays identified SlABI5-like7 and SlLEA2 as key transcriptional targets of SlWRKY37, integral to tomato seed dormancy regulation. Additionally, SlWRKY37 was found to be post-translationally phosphorylated at Ser65, a modification crucial for its transcriptional activation. Our findings elucidate the regulatory role of SlWRKY37 in seed dormancy, suggesting its potential as a target for gene editing to reduce seed dormancy in tomato breeding programs.

Methodological and Physiological Study during Seed Dormancy Release of Symplocos paniculata.

Symplocos paniculata are reported to exhibit seed dormancy, which impedes its cultivation and widespread adoption. In this study, a comprehensive method was established to overcome seed dormancy by subjecting seeds to scarification in 98% H2SO4 for 10 min, followed by 1000 mg·L-1 GA3 soaking for 48 h and stratification at 4 °C for 100 days. The seed germination percentage has increased significantly, to a peak of 42.67%, though the seeds could not germinate timely by NaOH scarification. Additionally, the dynamic changes of key stored substances (proteins, soluble sugars, starches, and fats), associated enzyme activities (amylases, peroxidase, and catalase), and endogenous hormones (abscisic acid, gibberellic acid, and indole-3-acetic acid) in seeds were investigated. The results demonstrated a continuous degradation of starch and fat in S. paniculata seeds, while the levels of protein and soluble sugar exhibited fluctuations, which probably facilitated seed dormancy breaking through energy supply and transformation. The enzymatic activities underwent rapid changes, accompanied by a gradual decrease in ABA content within the seeds with increasing stratification time. Notably, GA3, GA3/ABA, and (GA3 + IAA)/ABA showed significant increases, indicating their positive regulatory roles in seed germination. This study clarified the dormancy mechanism and established an effective method for the release dormancy of S. paniculata seeds.

Assessing Falling Number Stability Increases the Genomic Prediction Ability of Pre-Harvest Sprouting Resistance in Common Winter Wheat.

Pre-harvest sprouting (PHS) resistance is a complex trait, and many genes influencing the germination process of winter wheat have already been described. In the light of interannual climate variation, breeding for PHS resistance will remain mandatory for wheat breeders. Several tests and traits are used to assess PHS resistance, i.e., sprouting scores, germination index, and falling number (FN), but the variation of these traits is highly dependent on the weather conditions during field trials. Here, we present a method to assess falling number stability (FNS) employing an after-ripening period and the wetting of the kernels to improve trait variation and thus trait heritability. Different genome-based prediction scenarios within and across two subsequent seasons based on overall 400 breeding lines were applied to assess the predictive abilities of the different traits. Based on FNS, the genome-based prediction of the breeding values of wheat breeding material showed higher correlations across seasons (r=0.505-0.548) compared to those obtained for other traits for PHS assessment (r=0.216-0.501). By weighting PHS-associated quantitative trait loci (QTL) in the prediction model, the average predictive abilities for FNS increased from 0.585 to 0.648 within the season 2014/2015 and from 0.649 to 0.714 within the season 2015/2016. We found that markers in the Phs-A1 region on chromosome 4A had the highest effect on the predictive abilities for FNS, confirming the influence of this QTL in wheat breeding material, whereas the dwarfing genes Rht-B1 and Rht-D1 and the wheat-rye translocated chromosome T1RS.1BL exhibited effects, which are well-known, on FN per se exclusively.

Regulation of seed germination: ROS, epigenetic, and hormonal aspects.

BACKGROUND: The whole life of a plant is regulated by complex environmental or hormonal signaling networks that control genomic stability, environmental signal transduction, and gene expression affecting plant development and viability. Seed germination, responsible for the transformation from seed to seedling, is a key initiation step in plant growth and is controlled by unique physiological and biochemical processes. It is continuously modulated by various factors including epigenetic modifications, hormone transport, ROS signaling, and interaction among them. ROS showed versatile crucial functions in seed germination including various physiological oxidations to nucleic acid, protein, lipid, or chromatin in the cytoplasm, cell wall, and nucleus. AIM: of review: This review intends to provide novel insights into underlying mechanisms of seed germination especially associated with the ROS, and considers how these versatile regulatory mechanisms can be developed as useful tools for crop improvement. KEY SCIENTIFIC CONCEPTS OF REVIEW: We have summarized the generation and elimination of ROS during seed germination, with a specific focus on uncovering and understanding the mechanisms of seed germination at the level of phytohormones, ROS, and epigenetic switches, as well as the close connections between them. The findings exhibit that ROS plays multiple roles in regulating the ethylene, ABA, and GA homeostasis as well as the Ca2+ signaling, NO signaling, and MAPK cascade in seed germination via either the signal trigger or the oxidative modifier agent. Further, ROS shows the potential in the nuclear genome remodeling and some epigenetic modifiers function, although the detailed mechanisms are unclear in seed germination. We propose that ROS functions as a hub in the complex network regulating seed germination.

Long days induce adaptive secondary dormancy in the seeds of the Mediterranean plant Aethionema arabicum.

Secondary dormancy is an adaptive trait that increases reproductive success by aligning seed germination with permissive conditions for seedling establishment. Aethionema arabicum is an annual plant and member of the Brassicaceae that grows in environments characterized by hot and dry summers. Aethionema arabicum seeds may germinate in early spring when seedling establishment is permissible. We demonstrate that long-day light regimes induce secondary dormancy in the seeds of Aethionema arabicum (CYP accession), repressing germination in summer when seedling establishment is riskier. Characterization of mutants screened for defective secondary dormancy demonstrated that RGL2 mediates repression of genes involved in gibberellin (GA) signaling. Exposure to high temperature alleviates secondary dormancy, restoring germination potential. These data are consistent with the hypothesis that long-day-induced secondary dormancy and its alleviation by high temperatures may be part of an adaptive response limiting germination to conditions permissive for seedling establishment in spring and autumn.

Soil salinity regulates spatial-temporal heterogeneity of seed germination and seedbank persistence of an annual diaspore-trimorphic halophyte in northern China.

BACKGROUND AND AIMS: Seed heteromorphism is a plant strategy that an individual plant produces two or more distinct types of diaspores, which have diverse morphology, dispersal ability, ecological functions and different effects on plant life history traits. The aim of this study was to test the effects of seasonal soil salinity and burial depth on the dynamics of dormancy/germination and persistence/depletion of buried trimorphic diaspores of a desert annual halophyte Atriplex centralasiatica. METHODS: We investigated the effects of salinity and seasonal fluctuations of temperature on germination, recovery of germination and mortality of types A, B, C diaspores of A. centralasiatica in the laboratory and buried diaspores in situ at four soil salinities and three depths. Diaspores were collected monthly from the seedbank from December 2016 to November 2018, and the number of viable diaspores remaining (not depleted) and their germinability were determined. RESULTS: Non-dormant type A diaspores were depleted in the low salinity "window" in the first year. Dormant diaspore types B and C germinated to high percentages at 0.3 and 0.1 mol L-1 soil salinity, respectively. High salinity and shallow burial delayed depletion of diaspore types B and C. High salinity delayed depletion time of the three diaspore types and delayed dormancy release of types B and C diaspores from autumn to spring. Soil salinity modified the response of diaspores in the seedbank by delaying seed dormancy release in autum and winter and by providing a low-salt concentration window for germination of non-dormant diaspores in spring and early summer. CONCLUSIONS: Buried trimorphic diaspores of annual desert halophyte A. centralasiatica exhibited diverse dormancy/germination behavior in respond to seasonal soil salinity fluctuation. Prolonging persistence of the seedbank and delaying depletion of diaspores under salt stress in situ primarily is due to inhibition of dormancy-break. The differences in dormancy/germination and seed persistence in the soil seedbank may be a bet-hadging strategy adapted to stressful temporal and spatial heterogeneity, and allows A. centralasiatica to persist in the unpredictable cold desert enevironment.