This review provides a revised and expanded word-formula system of whole-seed primary dormancy classification that integrates the scheme of Nikolaeva with that of Baskin and Baskin. Notable changes include the following. (1) The number of named tiers (layers) in the classification hierarchy is increased from three to seven. (2) Formulae are provided for the known kinds of dormancy. (3) Seven subclasses of class morphological dormancy are designated: ‘dust seeds’ of mycoheterotrophs, holoparasites and autotrophs; diaspores of palms; and seeds with cryptogeal germination are new to the system. (4) Level non-deep physiological dormancy (PD) has been divided into two sublevels, each containing three types, and Type 6 is new to the system. (5) Subclass epicotyl PD with two levels, each with three types, has been added to class PD. (6) Level deep (regular) PD is divided into two types. (7) The simple and complex levels of class morphophysiological dormancy (MPD) have been expanded to 12 subclasses, 24 levels and 16 types. (8) Level non-deep simple epicotyl MPD with four types is added to the system. (9) Level deep simple regular epicotyl MPD is divided into four types. (10) Level deep simple double MPD is divided into two types. (11) Seeds with a water-impermeable seed coat in which the embryo-haustorium grows after germination (Canna) has been added to the class combinational dormancy. The hierarchical division of primary seed dormancy into many distinct categories highlights its great diversity and complexity at the whole-seed level, which can be expressed most accurately by dormancy formulae.

‘Dust seeds’ with an undifferentiated (organless) embryo are known to be produced by mycoheterotrophic species (MH) in nine families of angiosperms. However, aside from the numerous studies on seed germination of orchids, relatively little is known about germination in MH families. In the Ericaceae, some degree of mycoheterotrophy (full, partial or initial) and dust seeds with an undifferentiated embryo occur in all species in the three tribes of Monotropoideae, the only subfamily of Ericaceae with this combination of characters. In most species, the seed is <0.90 mm in the greatest dimension, the endosperm is absent (Pityopus) or consists of few to many (30–40) cells, and the embryo is minute, consisting of as few as two cells in Monotropa. Germination in Monotropoideae is monopolar, with only the radicular pole of the embryo participating in germination. Thus, germination polarity differs from that of the dust seeds of orchids in which only the plumular pole of the embryo (protocorm) participates in germination. The dust seeds in Monotropoideae require the presence of fungi, either direct contact with a fungus or the presence of a diffusible substance therefrom, to germinate (symbiotic germination). Recently, representatives of the four genera of tribe Pyroleae have been successfully germinated asymbiotically in vitro. We present a broad overview of dust-size seeds in angiosperms and conclude that they should be subdivided into at least two major categories.

Thermal time models have been widely applied to predict temperature requirements for seed germination. Generally, a log-normal distribution for thermal time [θT(g)] is used in such models at suboptimal temperatures to examine the variation in time to germination arising from variation in θT(g) within a seed population. Recently, additional distribution functions have been used in thermal time models to predict seed germination dynamics. However, the most suitable kind of the distribution function to use in thermal time models, especially at suboptimal temperatures, has not been determined. Five distributions (log-normal, Gumbel, logistic, Weibull and log-logistic) were used in thermal time models over a range of temperatures to fit the germination data for 15 species. The results showed that a more flexible model with the log-logistic distribution, rather than the log-normal distribution, provided the best explanation of θT(g) variation in 13 species at suboptimal temperatures. Thus, at least at suboptimal temperatures, the log-logistic distribution is an appropriate candidate among the five distributions used in this study. Therefore, the distribution of parameters [θT(g)] should be considered when using thermal time models to prevent large deviations; furthermore, an appropriate equation should be selected before using such a model to make predictions.

The genus Nolana (Solanaceae) comprises numerous species endemic to the coastal Atacama Desert of Chile and Peru of high ornamental potential and conservation value. The environments in which these species have evolved and are present today correspond to particular conditions in the midst of a hyper-arid habitat, so the study of their germination requirements and characterisation of seed dormancy becomes important in terms of conservation but also for ecological and evolutionary purposes. Different treatments were performed on mericarps of 12 species of Nolana: control (intact seeds imbibed in distilled water), scarification in funicular plug and distilled water and scarification in funicular plug and addition of GA3 (500 ppm); their permeability to water was also tested. It was determined that the species did not present physical dormancy, as had been previously reported, but rather physiological dormancy (PD). Germination results after treatments were not homogeneous among all 12 species, indicating differences in their dormancy levels. Also, the important role of the endosperm in the prevention of germination for the studied Nolana species was highlighted. Regarding the relationship between the level of PD (expressed as the percentage of germination for the most successful treatment) and the latitudinal distribution of the species or their phylogenetic closeness, it was determined that, for the studied species, their proximity in terms of clades was more relevant than their latitudinal distribution.

Fatty acid (FA) levels and profiles are vital for soybean oil quality, while cytokinins (CKs) and abscisic acid (ABA) are potent regulators of plant growth and development. Previous research suggested associations between FA biosynthesis and hormonal signalling networks; however, hormonal regulation of FA accumulation during soybean (Glycine max) seed maturation has never been measured. We analysed hormone and FA profiles obtained from HPLC-(ESI)-MS/MS and GC-FID screening during soybean seed maturation. A multilayered data processing approach, involving heat-maps, principal component analysis (PCA), correlation and multiregression models, suggested a strong relationship between hormone metabolism and FA/oil accumulation during seed maturation. Most strikingly, positive correlations were found between the levels of CK ribosides [transZeatin riboside (tZR), N6-isopentenyladenosine (iPR)] at the early stages of SM (R5-R6) and C18:0, C18:2 and oil content at the R8 stage. Moreover, multiple regression models revealed functional linkages between several CK derivatives and FA and oil content in mature seeds. To further test the significance of hormone regulation in FA metabolism, plants of two soybean accessions with contrasting hormone and FA profiles were sprayed with exogenous ABA and transZeatin (tZ) during the seed-filling period (R5-R6). Depending on the hormone type and concentration, these treatments distinctly modified biosynthesis of all tested FAs, except for C18:0. Most remarkably, tZ (50 nM) promoted production of C16:0, C18:1, C18:2, C18:3, and oil accumulation in maturing seeds. Overall, the results indicate impactful roles for ABA and CKs in FA accumulation during SM and represent a further step towards understanding FA biosynthesis, and potential improvements of soybean oil profiles.

Temperature may regulate seed dormancy and germination and determine the geographical distribution of species. The present study investigated the thermal limits for seed germination of Polygonum ferrugineum (Polygonaceae), an aquatic emergent herb distributed throughout tropical and subtropical America. Seed germination responses to light and temperature were evaluated both before (control) and after stratification at 10, 15 and 20°C for 7, 14 and 28 d. Germination of control seeds was ~50% at 10 and 15°C, and they did not germinate from 20 to 30°C. The best stratification treatment was 7 d at 10°C, where seed germination was >76% in the dark for all temperatures, except at 30°C, and < 60% in light conditions. A thermal time approach was applied to the seed germination results. Base temperature (Tb) was 6.3°C for non-dormant seeds and optimal temperature (To) was 20.6°C, ceiling temperature (Tc (<50)) was 32.8°C, and thermal time requirement for 50% germination was 44.4°Cd. We concluded that a fraction of P. ferrugineum seeds is dormant, has a narrow thermal niche to germinate (10 and 15°C) and that cold stratification (10°C) alleviated dormancy and amplified the thermal range permissive for germination of the species. Consequently, P. ferrugineum is expected to occur in colder environments, for example, at high altitudes. Higher temperatures decrease the probabilities of alleviate dormancy and the ability of their seeds to germinate.

For physiologically dormant (PD) species in fire-prone environments, dormancy can be both complex due to the interaction between fire and seasonal cues, and extremely deep due to long intervals between recruitment events. Due to this complexity, there are knowledge gaps particularly surrounding the dormancy depth and cues of long-lived perennial PD species. This can be problematic for both in situ and ex situ species management. We used germination experiments that tested seasonal temperature, smoke, dark and heat for 18 PD shrub species distributed across temperate fire-prone Australia and assessed how germination was correlated with environmental factors associated with their home environments. We found extremely high levels of dormancy, with only eight species germinating above 10% and three species producing no germination at all. Seven of these eight species had quite specific seasonal temperature requirements and/or very strong responses to smoke cues. The maximum germination for each species was positively correlated with the mean temperature of the source population but negatively correlated with rainfall seasonality and driest months. The strong dependence on a smoke cue for some of the study species, along with examples from other studies, provides evidence that an obligate smoke response could be a fire-adapted germination cue. Germination response correlated with rainfall season of the source populations is a pattern which has often been assumed but little comparative data across sites with different rainfall seasonality exists. Further investigation of a broader range of species from different rainfall season environments would help to elucidate this knowledge gap.

Although the level of seed desiccation sensitivity (LSDS) may have an impact on plant species conservation, information is available for <10% of tropical angiosperms. A study was conducted to assess the LSDS of 28 tropical montane species in Sri Lanka. Seeds were extracted from freshly collected fruits. Initial weight was recorded, and thousand seed weight (TSW) was calculated. Seed moisture content (MC) was determined. LSDS was determined using seed desiccation experiments and predicted using the TSW–MC criterion. Seed storage behaviour was predicted using LSDS and storage data and using a model based on phylogenetic affiliation. The relationship between LSDS and seed dormancy, life form and forest strata was evaluated. Fresh seeds of only 12 species germinated to >80%. Although seeds of the other species had >80% viability, only 0–70% germinated due to dormancy. Seeds of five species had MC <15%, indicating desiccation tolerance (DT). Seeds of 12 species lost viability after desiccation, indicating desiccation sensitivity (DS). Seeds of Ardisia missionis, Psychotria gartneri and Psychotria nigra remained viable after desiccation, showing DT. Seeds of 17 species were DS and those of 11 species DT. The TSW of four species was >500 g. Thus, seeds of other species were predicted to be DT by the TSW–MC criterion. A relationship was identified between LSDS and the forest strata of the species. More canopy species produced DS than DT seeds. Since seeds of most of the studied species were DS, these species may be threatened due to prolonged droughts predicted for the region due to climate change.

Genetic susceptibility to late maturity alpha-amylase (LMA) in wheat (Triticum aestivum L.) results in increased alpha-amylase activity in mature grain when cool conditions occur during late grain maturation. Farmers are forced to sell wheat grain with elevated alpha-amylase at a discount because it has an increased risk of poor end-product quality. This problem can result from either LMA or preharvest sprouting, grain germination on the mother plant when rain occurs before harvest. Whereas preharvest sprouting is a well-understood problem, little is known about the risk LMA poses to North American wheat crops. To examine this, LMA susceptibility was characterized in a panel of 251 North American hard spring wheat lines, representing ten geographical areas. It appears that there is substantial LMA susceptibility in North American wheat since only 27% of the lines showed reproducible LMA resistance following cold-induction experiments. A preliminary genome-wide association study detected six significant marker-trait associations. LMA in North American wheat may result from genetic mechanisms similar to those previously observed in Australian and International Maize and Wheat Improvement Center (CIMMYT) germplasm since two of the detected QTLs, QLMA.wsu.7B and QLMA.wsu.6B, co-localized with previously reported loci. The Reduced height (Rht) loci also influenced LMA. Elevated alpha-amylase levels were significantly associated with the presence of both wild-type and tall height, rht-B1a and rht-D1a, loci in both cold-treated and untreated samples.

Smoke-derived seed germination is an important trait for plants to colonize postfire habitats. The well-characterized smoke-derived chemicals of karrikins germinate seeds of species not known to occur after fires in nature. Hence, the ecologically relevant germination cues in smoke remain to be explored for native postfire plants. With the fire-chaser, Nicotiana attenuata, we revisit a bioassay-driven fractionation of liquid smoke to identify ecologically relevant germination cues. By combining bioassay-guided fractionation and comparative unbiased metabolomics, we developed a robust and efficient method to identify germination cues in smoke. Syringaldehyde (SAL) was re-identified as a germination cue in fractions of liquid smoke that promote seed germination. SAL was found to be produced during wildfires in the plant's native habitat, efficiently adsorbed to N. attenuata seeds from aqueous solutions and not readily leached from soil and accurately predicted the boundaries of natural fire events that reflect the occurrence of native postfire N. attenuata populations. We propose that SAL is an ecologically relevant germination cue in smoke for this species.

Functional traits related to regeneration responses to the environment are highly determinants of distribution patterns of plant communities. A large body of studies on seed traits suggests that regional climate may act as a strong filter of plant recruitment; however, few studies have evaluated the relative importance of seed traits and environmental filters for seed persistence at the population level. We tested the role of seed mass, water content and desiccation tolerance, as well as the germination time as proxies for seed tolerance to environmental filters (water deficit, heat shock and high temperatures) by comparing the response of tree species co-occurring in savannas located in different regions: Cerrado biome of Central Brazil and the Rio Branco savannas of northern Brazil. Seeds collected in savannas of Rio Branco showed a higher tolerance to environmental filters than those collected in savannas of the Cerrado. While the germination percentages largely varied in response to the treatments, the germination times were virtually unaffected by them, irrespective of seed origin, seed mass and water content. At the population level, the regional environment was a key determinant of seed tolerance to stress, irrespective of seed traits. Germination time was shown to represent a conservative seed trait and more linked to a species-specific germination strategy than to regional characteristics. Our results suggest that recruitment patterns of Cerrado savannas may be more impacted than Rio Branco savannas by the climate scenarios predicted for the future.

Luiz Fernando Gouvêa Labouriau (1921–1996) was a pioneer plant biologist who made significant contributions to plant physiology, mostly by bringing seed germination into a thermobiological context. His studies have set the foundations of seed science in Brazil and keep inspiring researchers until now.

The objective of this study was to determine how the current (10–16 weeks) and predicted future (2–8 weeks) length of cold stratification and current and predicted future post-stratification temperatures influence radicle and epicotyl emergence in acorns of Quercus robur. We tested radicle and epicotyl emergence at two temperatures corresponding to the current (15/6°C) and predicted future early autumn and spring temperatures (25/15°C) in Poland. We fitted models to describe and derive parameters for radicle and epicotyl emergences over time. The parameters included maximum percentage, rate of emergences, time to achieve the maximum emergence rate, emergence delay and time to 50% emergence. In most cases, the Gompertz model was the best fit, but in a few cases, the logistic model was the best. Richard's model for most of the cases did not converge. This model, according to both information criteria values, was the best fit for epicotyl emergence at 15/6°C following 8 weeks of cold stratification. Richard's model was also the best fit for epicotyl emergence at 25/15°C following 14 weeks of stratification.. Our results indicate that at temperatures typical for early autumn (15/6°C), the time necessary for radicle emergence from 50% of acorns was longer than that from acorns placed at 25/15°C. Four weeks of cold stratification extended 50% radicle emergence at 15/6°C to 70 d, whereas 12 weeks of stratification shortened the time to 11 d. When the acorns were incubated at 25/15°C, radicle emergence occurred faster than at 15/6°C and the time lag between radicle and epicotyl was shorter.

The wheat industry rejects grain with unacceptably high α-amylase enzyme levels due to the risk of poor endproduct quality. There are two main causes of elevated grain α-amylase: (1) preharvest sprouting in response to rain before harvest and (2) late maturity α-amylase (LMA) induction in response to a cool temperature shock during late grain development. LMA induction was detected in a panel of 24 Northwestern US spring wheat lines. Thus, this problem previously described in Australian and U.K. varieties also exists in U.S. varieties. Because LMA induction results were highly variable using published methods, a characterization of LMA-inducing conditions was conducted in an LMA-susceptible soft white spring wheat line, WA8124. Problems with elevated α-amylase in untreated controls were reduced by raising the temperature, 25°C day/18°C night versus 20°C day/10°C night. LMA induction was not improved by colder temperatures (15°C day/4°C night) versus moderately cold temperatures (18°C day/7.5°C night or 10°C day/10°C night). While previous studies observed LMA induction by heat stress, it failed to induce LMA in WA8124. Thus, not all LMA-susceptible cultivars respond to heat. The timing of LMA susceptibility varied between two cultivars and within a single cultivar grown at slightly different temperatures. Thus, variability in LMA induction likely results from variability in the timing of the grain developmental stage during which cold shock induces LMA. Thus, it was concluded that the visual inspection of grain is needed to correctly identify LMA-sensitive spikes at the soft dough stage of grain development (Zadok's stage 85).

Predispersal seed predation is one of the main causes of seed mortality in plant populations, contributing to decreased plant recruitment. Seed loss has previously been found to be related to crop size. Thus, we examined the influence of individual crop size on predispersal seed predation by beetles in the palm Syagrus flexuosa in the Brazilian savanna. The study was carried out in three tropical woodland savanna sites, where we sampled the total seed crop of 46 fruiting palms and checked the presence of beetle larvae inside all seeds per plant. We observed predispersal seed predation of S. flexuosa from all sites and a high variation in the number of seeds preyed on per individual palm. Crop size had a positive influence on the number of seeds lost to predispersal seed predators. Variations in levels of predispersal seed predation may also be accounted for by the reproductive phenology of S. flexuosa. If fruits are not available at the same time, less resource is available for predators and therefore a high proportion of seeds may be preyed on. Thus, our study demonstrates that an individual plant trait, crop size, is an important predictor of beetle seed damage per palm and a driver of the number of seeds lost to predispersal seed predators.

Large herbivores can disperse seeds over long distances through endozoochory. The Korean water deer (Hydropotes inermis argyropus), an internationally vulnerable species but locally considered a vermin, is a potential endozoochorous seed dispersal vector. In this study, feeding experiments were conducted to test the efficiency of seed dispersal through gut ingestion by the Korean water deer, its temporal pattern and the effect of gut passage on seed recovery and germination rate. Eight plant species, including species that formerly germinated from its faeces, were used to feed three Korean water deer. Once the deer had consumed all the provided seeds, their faeces were collected after 24, 48, 72 and 96 h. The collected faeces were air-dried, and the number of seeds retrieved from the faeces was counted every 24 h (0–24, 24–48, 48–72 and 72–96 h). Among the eight plant species, six species were retrieved with intact seeds. Panicum bisulcatum had the highest recovery rate of 33.7%, followed by Amaranthus mangostanus (24.5%) and Chenopodium album (14.4%). Most of the seeds were recovered within the 24–48 h time interval. Germination tests were conducted on the ingested and uningested seeds for the four species which had a sufficient recovery rate. The effects of gut passage on seed germination differed according to plant species. The germination rate substantially decreased after gut passage. The results suggest that the Korean water deer can disperse seeds, potentially over long distances albeit at a high cost of low seed recovery and germination rate.

Pre-harvest sprouting (PHS) is a common threat to cereal crops in which the grain maturation phase takes place under rainy, moist conditions. Susceptibility to PHS is higher in sorghum genotypes displaying low levels of seed dormancy before harvest maturity. Other attributes such as glume or panicle morphology may also affect susceptibility to PHS. Breeding for resistance to PHS in grain sorghum requires the identification of grain physiological and morphological attributes affecting this trait, and a protocol for phenotyping and rating genotypes according to their susceptibility to PHS. In this work, we tested germination under laboratory conditions using detached grains and intact panicles for a panel of 20 sorghum genotypes including 11 parental lines, 6 hybrids and 3 reference inbred lines with contrasting PHS response. Records for natural sprouting in the field for these genotypes were also included in the analysis. Multivariate analyses of germination data allowed separation of genotypes into two major categories (resistant and susceptible to PHS). Laboratory germination data correlated significantly with PHS in the field. In most genotypes, the glumes had a significant, inhibitory effect on germination. The low levels of grain dormancy were observed among high tannin backgrounds, and vice versa, indicating that a pigmented testa alone does not provide resistance to PHS. Altogether, the phenotyping protocol allowed the classification of sorghum genotypes according to their susceptibility to PHS and the identification of different attributes useful for breeding for PHS resistance in this crop.

A low-cost technique named ‘on-farm’ seed priming is increasingly being recognized as an effective approach to maximize crop establishment. It consists of anaerobically soaking seeds in water before sowing resulting in rapid and uniform germination, and enhanced seedling vigour. The extent of these benefits depends on the soaking time. The current determination of optimal soaking time by germination assays and mini-plot trials is resource-intensive, as it is species/genotype-specific. This study aimed to determine the potential of the seed respiration rate (an indicator of metabolic activity) and seed morphological changes during barley priming as predictors of the priming benefits and, thus, facilitate the determination of optimal soaking times. A series of germination tests revealed that the germination rate is mostly attributable to the rapid hydration of embryo tissues, as the highest gains in the germination rate occurred before the resumption of respiration. Germination uniformity, however, was not significantly improved until seeds were primed for at least 8 h, that is, after a first respiration burst was initiated. The maximum seedling vigour was attained when the priming was stopped just before the beginning of the differentiation of embryonic axes (20 h) after which vigour began to decrease (‘over-priming’). The onset of embryonic axis elongation was preceded by a second respiration burst, which can be used as a marker for priming optimization. Thus, monitoring of seed respiration provides a rapid and inexpensive alternative to the current practice. The method could be carried out by agricultural institutions to provide recommended optimal soaking times for the common barley varieties within a specific region.