Seed enhancements may be defined as post-harvest treatments that improve germination or seedling growth, or facilitate the delivery of seeds and other materials required at the time of sowing. This definition includes three general areas of enhancements: pre-sowing hydration treatments (priming), coating technologies and seed conditioning. Pre-sowing hydration treatments include non-controlled water uptake systems (methods in which water is freely available and not restricted by the environment) and controlled systems (methods that regulate seed moisture content preventing the completion of germination). Three techniques are used for controlled water uptake: priming with solutions or with solid particulate systems or by controlled hydration with water. These priming techniques will be discussed in this paper with reference to methodology, protocol optimization, drying and storage. Coating technologies include pelleting and film coating, and coatings may serve as delivery systems. Seed conditioning equipment upgrades seed quality by physical criteria. Integration of these methods can be performed, and a system is described to upgrade seed quality in Brassica that combines hydration, coating and conditioning. Upgrading is achieved by detecting sinapine leakage from nonviable seeds in a coating material surrounding the seeds. Seed-coat permeability directly influences leakage rate, and seeds of many species have a semipermeable layer. The semipermeable layer restricts solute diffusion through the seed coat, while water movement is not impeded. Opportunities for future seed enhancement research and development are highlighted.

Bromus tectorum L. is an invasive winter annual grass with seeds that lose dormancy through the process of dry after-ripening. This paper proposes a model for after-ripening of B. tectorum seeds based on the concept of hydrothermal time. Seed germination time course curves are modelled using five parameters: a hydrothermal time constant, the fraction of viable seeds in the population, base temperature, mean base water potential and the standard deviation of base water potentials in the population. It is considered that only mean base water potential varies as a function of storage duration and incubation temperature following after-ripening. All other parameters are held constant throughout after-ripening and at all incubation temperatures. Data for model development are from seed germination studies carried out at four water potentials (0, −0.5, −1.0 and −1.5 MPa) at each of two constant incubation temperatures (15 and 25°C) following different storage intervals including recently harvested, partially after-ripened (stored for 4, 9 or 16 weeks at 20°C) and fully after-ripened (stored for 14 weeks at 40°C). The model was fitted using a repeated probit regression method, and for the two seed populations studied gave R2 values of 0.898 and 0.829. Germination time course curves predicted by the model generally had a good fit when compared with observed curves at the incubation temperature/water potential treatment combinations for different after-ripening intervals. Changes in germination time course curves during after-ripening of B. tectorum can largely be explained by decreases in the mean base water potential. The simplicity and good fit of the model give it considerable potential for extension to simulation of after-ripening under field conditions.

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After-ripening, the loss of dormancy under dry conditions, is associated with a decrease in mean base water potential for germination ofBromus tectorumL. seeds. After-ripening rate is a linear function of temperature above a base temperature, so that dormancy loss can be quantified using a thermal after-ripening time (TAR) model. To incorporate storage water potential into TAR, we created a hydrothermal after-ripening time (HTAR) model. Seeds from twoB. tectorumpopulations were stored under controlled temperatures (20 or 30 °C) and water potentials (−400 to −40 MPa). Subsamples were periodically removed from each storage treatment and incubated at 15 or 25 °C to determine germination time courses. Dormancy status (mean base water potential) was calculated from each time course using hydrothermal time equations developed for each seed collection. Seeds stored at −400 MPa did not after-ripen. At water potentials from −400 to −150 MPa, the rate of after-ripening increased approximately linearly with increasing water potential. Between −150 and −80 MPa, there was no further increase in after-ripening rate, while at −40 MPa seeds did not after-ripen and showed loss of vigour. These results suggest that the concept of critical water potential thresholds, previously shown to be associated with metabolic activity and desiccation damage in partially hydrated seeds, is also relevant to the process of after-ripening. The HTAR model generally improved field predictions of dormancy loss when the soil was very dry. Reduced after-ripening rate under such conditions provides an ecologically relevant explanation of how seeds prolong dormancy at high summer soil temperatures.

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Hydrothermal time (HTT) analysis is an effective method for quantifying germination response to temperature and water potential. Time course curves under any temperature and water potential conditions can be generated from knowledge of the HTT parameters mean base water potential, standard deviation of base water potentials, base temperature, and hydrothermal time requirement. Here we use these parameters as indices for making comparisons among 24 species including shrubs, grasses and perennial herbs from the deserts and semi-deserts of Asia and North America. Halophytes (salt-tolerant species) are characterized by low base water potential values but high hydrothermal time constants. Psammophytes (species that inhabit high-sand soils) are just the opposite, and bodenvags (generalist species with no special soil requirements) display a wide range in all parameter values. Variation in the distribution of base water potentials strongly influences uniformity of germination. The results illustrate that germination rate in water or at reduced water potentials is closely associated with HTT parameters. These findings also have important ecological relevance, in that they help explain differences in germination patterns associated with contrasting habitats.

Advances in seed biology include progress in understanding the ecological significance of seed dormancy mechanisms. This knowledge is being used to make more accurate predictions of germination timing in the field. For several wild species whose seedlings establish in spring, seed populations show relevant variation that can be correlated with habitat conditions. Populations from severe winter sites, where the major risk to seedlings is frost, tend to have long chilling requirements or to germinate very slowly at low temperatures. Populations from warmer sites, where the major risk is drought, are non-dormant and germinate very rapidly under these same conditions. Seed populations from intermediate sites exhibit variation in dormancy levels, both among and within plants, which spreads germination across a considerable time period. For grasses that undergo dry after-ripening, seed dormancy loss can be successfully modelled using hydrothermal time. Dormancy loss for a seed population is associated with a progressive downward shift in the mean base water potential, i.e., the water potential below which half of the seeds will not germinate. Other parameters (hydrothermal time requirement, base temperature and standard deviation of base water potentials) tend to be constant through time. Simulation models for predicting dormancy loss in the field can be created by combining measurements of seed zone temperatures with equations that describe changes in mean base water potential as a function of temperature. Successful validation of these and other models demonstrates that equations based on laboratory data can be used to predict dormancy loss under widely fluctuating field conditions. Future progress may allow prediction of germination timing based on knowledge of intrinsic dormancy characteristics of a seed population and long-term weather patterns in the field.

Hydrothermal time (HTT) describes progress toward seed germination under various combinations of incubation water potential ( ) and temperature (T). To examine changes in HTT parameters during dormancy loss, seeds from two populations of the bunchgrass Elymus elymoides were incubated under seven temperature regimes following dry storage at 10, 20 and 30°C for intervals from 0 to 16 weeks. Fully after-ripened seeds were primed for 1 week at a range of s. Data on germination rate during priming were used to obtain a HTT equation for each seed population, while data obtained following transfer to water were used to calculate HTT accumulation during priming. HTT equations accurately predicted germination time course curves if mean base water potential, b(50), was allowed to vary with temperature. b(50) values increased linearly with temperature, explaining why germination rate does not increase with temperature in this species. b(50) showed a linear decrease as a function of thermal time in storage. Slopes for the T × b(50) relationship did not change during after-ripening. This thermal after-ripening time model was characterized by a single base temperature and a constant slope across temperatures for each collection. Because the difference between initial and final b(50)s was uniform across tempera-tures, the thermal after-ripening requirement was also a constant. When seeds were primed for 1 week at −4 to −20 MPa, accumulation of HTT was a uniform 20% of the total HTT requirement. When primed at 0 to −4 MPa, HTT accumulation decreased linearly with decreasing priming potential, and a hydrothermal priming time model using a constant minimum priming potential adequately described priming effects. Use of these simple HTT relationships will facilitate modelling of germination phenology in the field.