A review is presented of work carried out over the last 40 years on non-porous sorbents and on microporous shape-selective sorbents derived from palygorskite, smectite, and vermiculite. Among such materials four kinds of behavior were observed. In some systems uptake was restricted to external surfaces; in others, intercalation also occurred but only above threshold pressures. If the interlayer region was completely filled by long-chain organic cations, imbibition was possible, but in amounts which were very strongly dependent upon the cohesive energy density of the sorbate. Finally, in certain permanently expanded derivatives of layer silicates intercalation proceeded without any threshold pressure, just as in zeolites.

In this latter group, micropores existed which sometimes resulted in shape-selective sorption and molecule sieving. The micropores in clay minerals were modified by varying the size and shape of the interlayer cations, their charge, and the charge density of the siliceous layers. The interplay of these factors was investigated and the micropore sorbents were shown to be highly effective in the separation of mixtures.

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.

The wheat bZip transcription factor TaABF1 mediates both abscisic acid (ABA)-induced and ABA-suppressed gene expression. As levels of TaABF1 protein do not change in response to ABA, and TaABF1 is in a phosphorylated state in vivo, we investigated whether TaABF1 could be regulated at the post-translational level. In bombarded aleurone cells, a TaABF1 protein carrying phosphomimetic mutations (serine to aspartate) at four sites (S36D, S37D, S113D, S115D) was three to five times more potent than wild-type TaABF1 in activating HVA1, an ABA-responsive gene. The phosphomimetic mutations also increased the ability of TaABF1 to downregulate the ABA-suppressed gene Amy32b. These findings strongly suggest that phosphorylation at these sites increases the transcriptional regulatory activity of TaABF1. In contrast to the activation observed by the quadruple serine to aspartate mutation, a single S113D mutation completely eliminated the ability of TaABF1 to upregulate HVA1 or downregulate Amy32b. Thus phosphorylation of TaABF1 can either stimulate or inhibit the activity of TaABF1 in regulating downstream genes, depending on the site and pattern of phosphorylation. Mutation of S318 and S322 (in the bZIP domain) eliminated the ability of TaABF1 to activate HVA1, but had no effect on the ability of TaABF1 to downregulate Amy32b, suggesting that TaABF1 represses Amy32b expression through a mechanism other than direct DNA binding. An important step towards understanding how ABA and gibberellin (GA) signals are integrated through TaABF1 phosphorylation to regulate downstream gene expression is to clarify the effects of those hormones on the expression of specific genes. In contrast to some other ABA-induced genes, we found that HVA1 induction by ABA or TaABF1 is not inhibited by GA.

Physical dormancy (PY) occurs in at least 18 angiosperm plant families and is caused by water-impermeable palisade cells in seed (or fruit) coats. Breaking of PY involves disruption or dislodgement of water-gap structures causing the seeds/fruits to become water permeable (non-dormant). The water-gap region is a morphologically distinct area of the seed or fruit coat that forms a water-gap complex. The location, anatomy, morphology and origin of water-gaps can differ between and even within families and genera. Water-gap structures sense environmental conditions that allow seeds with PY to become permeable just prior to the commencement of conditions favourable for germination and plant establishment. There are three basic water-gap morpho-anatomies characterized by the way the water-gap opens: Type-I, Type-II and Type-III. In Type-I water-gaps, specific kinds of cells pull apart to form a surface opening, while in Type-II a portion of the surface structure is pulled away from adjacent cells, opening the water-gap. Type-III is the least common type and has a circular, plug-like structure that is dislodged, whereby water entry occurs. In addition, water-gap complexes are either simple or compound, depending on whether only a single primary water-gap structure is involved in dormancy release or an additional secondary water-gap structure opens, permitting water entry.

Experiments were conducted to evaluate the influence of temperature and water potential on water uptake, germination, and emergence of large crabgrass in order to predict emergence in the field. Water uptake of seed soaked in polyethylene glycol solutions of 0 to −1400 kPa underwent an initial imbibition phase followed by a lag phase and subsequent increase in water content when radicles emerged from the seed. Maximum germination at 15 C was 12% at 0 kPa and 60% at 25 C at 0 to −200 kPa osmotic potential. In the growth chamber, large crabgrass emergence from soil began 2 to 3 d after planting at 30 or 35 C and within 9 to 10 d at 15 C. Maximum emergence of 77 % occurred at 25 C and at a soil water potential of −30 kPa. Emergence percentage decreased as water potential decreased or as temperature increased or decreased. A logistic equation described emergence of large crabgrass at each combination of temperature and soil water potential at which emergence occurred, and a predictive model was developed and validated by field data. In the field, there was little or no emergence at soil temperatures below 15 C or water potentials below −50 to −60 kPa. The model predicted the time of onset of large crabgrass emergence and the time to reach maximum emergence to within 2 to 4 d of that recorded in field experiments. The model also predicted the correct number of flushes of emergence occurring in the field in three of four experiments.

Laboratory studies were conducted to determine the effect of seed coat on woolly cupgrass seed moisture and oxygen uptake, and to determine if water-soluble growth inhibitors are present in the seed. Intact dormant seeds did not respond to any temperature regime or to oxygen concentrations above atmospheric level. Dehulling increased germination of dormant seed to about 85%. Oxygen concentrations above atmospheric level increased germination of partially dehulled (1/4 distal end of the seed dehulled) seeds an additional 10%. Oxygen uptake by dehulled dormant and intact nondormant seeds was greater than intact dormant seeds. Leaching intact dormant seeds did not promote germination, nor did the leachate inhibit germination of nondormant seeds. Embryos excised from dormant seeds germinated under laboratory conditions. Results from this study suggest that the woolly cupgrass seed coat may inhibit germination by controlling oxygen availability to the embryo.

The effect of exposing tame oat (Avena sativa L. ‘Hudson’) to trifluralin (α,α,α-trifluoro-2,6-dinitro-N,N-dipropyl-p-toluidine) and triallate [S-(2,3,3-trichloroallyl) diisopropylthiocarbamate] during seed imbibition was investigated in a growth cabinet at 21 C in the absence of light. Caryopses were imbibed in 0.0, 1.5, and 3.0 ppmv aqueous dilutions of trifluralin and triallate for up to 10 h, after which germination was continued in distilled water. Germination was not affected following imbibition in either herbicide. Both herbicides produced distinct visual symptoms of herbicide injury during subsequent seedling growth. Coleoptile length of 5-day-old seedlings was reduced following imbibition in a 1.5-ppmv dilution of either herbicide. Triallate at 1.5 ppmv resulted in a reduction in shoot dry weight. Both herbicides were largely excluded from embryos of imbibing caryopses. Seedlings were unaffected when cultured from embryos excised from caryopses imbibed in trifluralin and triallate dilutions. Seedling phytotoxic effects resulted from residual activity of trifluralin and triallate retained by the pericarp and testa of imbibed caryopses. Presence of the hull during imbibition decreased the phytotoxic effect of both herbicides on shoot growth. These findings suggest that germinating seedlings of Avena spp. may be affected by trifluralin and triallate at an earlier stage than previously realized.

Seeds (achenes) of fringed sage were collected in central Saskatchewan in 1987, 1990, and 1991, and studies were conducted to determine the effects of imbibition time before brief or prolonged exposure to red (R) and far-red (FR) light on germination. A majority of the seeds required light for germination, but a substantial proportion germinated in darkness. Phytochrome controlled germination, and its activity was influenced by the duration of imbibition and by seed collections. At least 8 h of imbibition were required before phytochrome was activated by R light. The stimulation of germination could be reversed by FR light, and this suppression subsequently could be reversed by exposure to R light. Variable light requirements for germination of fringed sage can be met following large- or small-scale disturbances or in relatively undisturbed conditions.

Increased osmotic potential from 0 to −14 bars decreased the moisture uptake and germination of mechanically scarified hemp sesbania [Sesbania exaltata (Raf.) Cory] seed. Germination percentage for osmotic potential of 0, −2, −4, −6, and −8 bars was 86%, 86%, 70%, 19%, and 1%, respectively. This response to moisture stress was modified by imbibition, or hydration-dehydration, of the seed prior to the moisture stress. Imbibing the seed for as little as 2 h significantly increased its subsequent germination against moisture stress. One hydration-dehydration cycle had little or no effect on germination, but as the number of cycles increased, the germination percentage decreased. Prolonged hydration of 3 and 7 days followed by dehydration greatly reduced subsequent seed germination.

Mexican palo-verde is a serious woody weed in tropical parts of the world. Like many such leguminous species, it has relatively large seeds with hard-seeded (physical) dormancy. It therefore has the potential for long-lived seed banks that are difficult to manage. The physiology of hard-seeded dormancy is still relatively poorly understood but has important implications for weed management. We propose that wet heat is a potentially important dormancy release mechanism for summer rainfall tropical regions. We described the relationships between wet heat and dormancy release (in water; three seed sources) and germination (near saturation; single seed source) by testing seeds at constant temperatures between 10 and 60 C. The logistic transformation of the temperature–dormancy relationship was best described by a quadratic equation below a threshold of ∼ 33.6 C and a linear equation above that threshold. The relationship was the same for all seed sources other than a phase shift of up to 6.6 C, which is likely to be of biological significance. Germination occurred between 15 and 40 C and was limited by cold stress at ≤ 20 C and heat stress > 35 C. The sensitivity of dormancy to naturally encountered temperature ranges suggests that wet heat is an important dormancy release mechanism and one that can be exploited when developing management strategies for invasive populations.

Glycine max cultivars exhibit differential tolerance to soil-applied sulfentrazone. The intent of this study was to determine the physiological basis for this differential tolerance by evaluating sulfentrazone absorption and metabolism during the earliest stages of G. max development (i.e., germinating seeds, and germinal seedlings). Imbibed seeds (24 h) of the sulfentrazone-tolerant cultivar ‘Stonewall’ absorbed 37% less sulfentrazone than the sulfentrazone-sensitive cultivar ‘Asgrow 6785’. Similarly, germinal seedlings (i.e., 60 h from start of imbibition) of the sulfentrazone-tolerant cultivars Stonewall and ‘Pioneer 9593’ absorbed 22% less sulfentrazone than the sulfentrazone-sensitive cultivars Asgrow 6785 and ‘Carver’ when exposed to sulfentrazone-containing solution for either 24 or 48 h. The amount of root-absorbed 14C-sulfentrazone that was translocated into cotyledon or hypocotyl tissues did not exceed 11% of the amount absorbed and was similar for all four cultivars. Sulfentrazone metabolism by both imbibed seeds and by germinal seedlings was independent of cultivar. Increasing the sulfentrazone concentration in the seed imbibition solution and increasing the temperature resulted in greater seedling height reduction at 10 d in Asgrow 6758 than in Stonewall. Results indicate that differential absorption during the earliest stages of development is the basis for the differential response among G. max cultivars. Comparatively limited sulfentrazone absorption by Stonewall, as reflected in acceptable seedling injury, remained relatively consistent across the range of concentrations and temperatures evaluated.

Leafy spurge seeds are responsive to alternating temperature rather than constant temperature for germination. Transcriptome changes of dry leafy spurge seeds and seeds imbibed for 1 and 3 d at 20 C constant (C) and 20 : 30 C alternating (A) temperature were determined by microarray analysis to examine temperature responsiveness. Principal component analysis revealed differences in the transcriptome of imbibed seeds based on the temperature regime. Computational methods in bioinformatics parsed the data into overrepresented AraCyc pathways and gene regulation subnetworks providing biological context to temperature responses. After 1 d of imbibition, the degradation of starch and sucrose leading to anaerobic respiration were common pathways at both temperature regimes. Several overrepresented pathways unique to 1 d A were associated with generation of energy, reducing power, and carbon substrates; several of these pathways remained overrepresented and up-regulated at 3 d A. At 1 d C, pathways for the phytohormones jasmonic acid and brassinosteroids were uniquely overrepresented. There was little similarity in overrepresented pathways at 1 d C between leafy spurge and arabidopsis seeds, indicating species-specific effects upon imbibition of dry seeds. Overrepresented gene subnetworks at 1 d and 3 d at both temperature regimes related to signaling processes and stress responses. A major overrepresented subnetwork unique to 1 d C related to photomorphogenesis via the E3 ubiquitin ligase COP1. At 1 d A, major overrepresented subnetworks involved circadian rhythm via LHY and TOC1 proteins and expression of stress-related genes such as DREB1A, which is subject to circadian regulation. Collectively, substantial differences were observed in the transcriptome of leafy spurge seeds imbibed under conditions that affect the capacity to germinate.

Seed mass is an important plant functional trait linked to germination. For instance, higher-mass seeds often display greater germination compared to lower-mass seeds when exposed to non-stressful conditions. Yet, knowledge pertaining to germination dynamics for different mass-based seed fractions following exposure to abiotic stress is lacking. Here, we assess the germination response of relatively fresh, mass-separated Rudbeckia mollis (Asteraceae) seeds to various simulated seasonal temperatures, supra-optimal temperatures and increasing ageing stress duration. Air density separation yielded three mass-based classes, called light (393 ± 35 μg), intermediate (423 ± 29 μg) and heavy (474 ± 38 μg). Water uptake kinetics indicated that imbibition (0–6 h) and germination lag (6–24 h) were independent of seed mass. Similarly, germination and viability loss of fresh seeds following exposure to seasonal and supra-optimal constant temperatures were independent of mass. However, seed mass influenced germination following increasing ageing stress, with light seeds germinating to a significantly greater extent than intermediate or heavy seeds. For example, final germination per cent in light-class seeds was about 1.7 times greater than intermediate or heavy seeds after 20 d of saturated salt accelerated ageing (SSAA). Seeds stored for 1 year in the laboratory displayed mass-dependent germination patterns similar to seeds following SSAA. Mass-independent germination responses may be a strategy to maintain an annual life history in otherwise difficult environments when R. mollis seeds are relatively fresh. However, differences in germination response between aged and unaged seeds suggest that mass-dependent viability loss may occur in R. mollis.

Glebionis coronaria (L.) Cass. ex Spach is a common Mediterranean weed producing distinctive central and peripheral dormant cypselae with a hard fruit coat, which was previously hypothesized to impose physical dormancy. Analysis of water uptake in cypselae and in naked seeds showed that it preferentially takes place at the basal end of the fruit; however, seeds within an intact pericarp do not fully imbibe when compared with naked seeds. Germination was not significantly different between the two heteromorphs, and afterripening or cold stratification did not increase germination, while warm stratification at 35/20°C, as revealed by logistic regression, resulted in a significant improvement. However, loss of viability was also rapid at these high temperatures. Central and peripheral cypselae generally showed very low germination. In both heteromorphs, faster and higher germination (60–70%) was reached only after extensive scarification of pericarp tissue, and full germination was observed only after complete removal of pericarp tissue. Although the pericarp significantly reduced water uptake, no palisade layer(s) of macrosclereids could be observed. Xylem-vessel elements were found running through the basal end of the pericarp and forming the main point of water entry. We reject the hypothesis that G. coronaria cypselae have physical dormancy. Instead, water uptake and germination are impeded by: (1) directed water uptake, mainly through a pericarp-spanning channel-like structure; and (2) mechanical constraint on embryo growth exerted by the hard pericarp. The channel-like structure forms the principal system for controlling seed germination.

Structural studies in seeds with physical dormancy (PY) are important to better understand its causes and release when subjected to treatments for dormancy breaking. The aims of this study were to (1) characterize the PY break; (2) examine the role of different seed structures in water uptake; and (3) identify the water gap in Senna multijuga seeds. Imbibition patterns of dormant and non-dormant (subjected to dormancy breaking treatments) seeds and the morphological changes during dormancy breaking and germination were evaluated. To identify the water gap, the micropyle and lens were blocked separately, and the water absorption by seed parts was determined. Structural characteristics of the seed coat were also examined. Immersion in water at 80°C was efficient in breaking seed dormancy and imbibition occurred first at the hilar region, through the lens. Water was not absorbed through the micropyle or the extra-hilar region. S. multijuga seeds have a testa with a linearly aligned micropyle, hilum and lens. The seed coat consisted of a cuticle, macrosclereids, one (hilar region) or two (extra-hilar region) layer(s) of osteosclereids and parenchyma cell layers. The lens has typical parenchyma cells underneath it and two fragile regions comprised of shorter macrosclereids. Heat treatment stimulated the lens region, resulting in the opening of fragile regions at the lens, allowing water to enter the seeds. It is concluded that short-term exposure to a hot water treatment is sufficient for the formation of a water gap in S. multijuga seeds, and only the lens acts in the imbibition process.

Seed germination is regulated in a concerted manner that involves generating growth potential in the embryo to overcome the mechanical resistance of the endosperm. The wake-up call of a dry seed includes the reorganization of subcellular structures and the reactivation of metabolism in a dense, oxygen-poor environment. Pools of unbound metabolites and solutes produced by the degradation of storage reserves, including starch, proteins and oils, in the embryo can contribute to the generation of the embryo growth potential and radicle protrusion. Recent genomics studies have contributed a vast amount of data on protein, metabolite and gene transcript profiles during germination, which can be integrated to explore the seed metabolism from water imbibition to radicle protrusion. To what extent are free pools of metabolites relevant to the reorganization of seed metabolism? How is energy built to support embryo growth and radicle protrusion? Elucidating these fundamental questions in seed biology is the key to the understanding of the germination process. Here we have attempted to summarize the recent scientific knowledge to provide a comprehensive description of the ignition, reassembling and regulation of metabolism during seed germination.

Physical seed dormancy is a common attribute among plants, and a wide range of dormancy-release mechanisms have been described, but their ecological significance is rarely tested through comparative study. This study tests whether dormancy-release responses to wet heat in four legume species with physical dormancy are correlated with habitat: two wetland species (Mimosa pigra and Parkinsonia aculeata, both dispersed primarily by water) and two terrestrial species (Acacia nilotica and Prosopis pallida, both dispersed primarily through vertebrate herbivores). Dormancy release was compared at three moisture levels (80% relative humidity, saturated and submerged) at constant (20–45°C) and diurnally fluctuating (20/40°C) temperatures for 14 d. Seed viability was tested by germinating at 25°C. The functional relationship between temperature and dormancy release after 14 d differed between species: submerged seeds of the two wetland species showed a quadratic response, with low rates of imbibition below 20–25°C and complete imbibition at around 40°C; P. pallida seeds showed a linear positive relationship, whereas there was no temperature response for A. nilotica seeds below 45°C. Surprisingly, dormancy release after 14 d was relatively insensitive to moisture levels, although rate of dormancy release was generally slower under drier conditions. Dormancy release was not influenced by fluctuating temperatures. Seed viability was largely unaffected by temperature or moisture regime, although it did differ with species and was lower for non-dormant seeds. Our results suggest that a functional dormancy-release response to wet heat provides important fitness benefits for wetland species, but not for species dispersed through vertebrate herbivores, for which it may be maladaptive.

The wheat transcription factor TaABF1 physically interacts with the protein kinase PKABA1 and mediates both abscisic acid (ABA)-induced and ABA-suppressed gene expression. In bombarded aleurone cells of imbibing grains, the effect of TaABF1 in down-regulating the gibberellin (GA)-induced Amy32b promoter was stronger in the presence of exogenous ABA. As these grains contained low levels of endogenous ABA, the effect of TaABF1 may also be mediated by ABA-induced activation even in the absence of exogenous ABA. Levels of TaABF1 protein decreased slightly during imbibition of afterripened grains. However, TaABF1 levels (especially in aleurone layers) were not substantially affected by exogenous ABA or GA, indicating that changes in TaABF1 protein level are not an important part of regulating its role in hormone signalling. We found that TaABF1 was phosphorylated in vivo in aleurone cells, suggesting a role for post-translational modification in regulating TaABF1 activity. Induction of Amy32b by overexpression of the transcription factor GAMyb could not be prevented by TaABF1, indicating that TaABF1 acts upstream of GAMyb transcription in the signalling pathway. Supporting this view, knockdown of TaABF1 by RNA interference resulted in increased expression from the GAMyb promoter. These results are consistent with a model in which TaABF1 is constitutively present in aleurone cells, while its ability to down-regulate GAMyb is regulated in response to ABA.

Morphological changes in tomato (Lycopersicon esculentum Mill. cv. Moneymaker) seed during osmopriming and imbibition (‘hydropriming’) were followed using X-ray photographs. Embryo, endosperm and free space areas were measured. Both osmopriming and hydropriming resulted in free space development (+ 8.1% and + 10.8% of the whole seed planar area, respectively), almost all at the cost of the endosperm area. Planar dimensions of whole seeds were relatively constant and the dimension perpendicular to the planar surface, the thickness, could account for the volume increase of primed seeds reported in the literature. In dead seeds, only a small amount of free space developed while the planar area of the seed remained the same. In the imbibing viable seeds no deterioration of endosperm could be detected until the moment of root protrusion.

Seeds which were osmoprimed directly after harvest, i.e. in the fresh state, did not showthe induction of any free space, while free space developed normally after dehydration and a second priming treatment. Apparently, a dehydration step prior to the priming treatments is required for the development of free space in osmo- or hydroprimed seed.

X-ray photographs of hydroprimed seeds showed that the radicle tip adhered tightly to theendosperm cap. This results in various forms of damage to the root tips upon redrying. Priming did not introduce cotyledon abnormalities.

Inorganic P, total P and phytic acid-P leakage from imbibing whole soybean, bean, and pea seeds, as well as whole maize and barley grains were measured. The amounts of P and phytic acid leaked during a 6-h soaking period varied for the five species. Soybeans leaked the most P and phytic acid of the species examined. Peas, beans and barley leaked little or no phytic acid. Based on reported levels of total P and phytic acid in the dry seeds/grains that were studied, total P leakage ranged from 0.5 to 1.2%, while phytic acid leakage ranged from 0.01 to 0.5%. No relationship was found between initial levels of total P or phytic acid in the dry seeds/grains and the amount of total P or phytic acid leaked by the imbibed seed/grain. Similarly, the amounts of phytic acid leaked by the seeds/grains of this study did not appear to be related to the water solubility of the phytic acid in the dry seeds/grains.