Rapid seed germination and stand establishment are critical factors to crop production under salt‐stress conditions. In many crop species, seed germination and early seedling growth are the most sensitive stages to salinity stress. Salinity may delay the onset, reduce the rate, and increase the dispersion of germination events, leading to reductions in plant growth and final crop yield. The adverse effects of salt‐stress can be alleviated by various measures, including seed priming (a.k.a. pre‐sowing seed treatment). The general purpose of seed priming is to partially hydrate the seed to a point where germination processes are begun but not completed. Most priming treatments involve imbibing seed with restricted amounts of water to allow sufficient hydration and advancement of metabolic processes but preventing germination or loss of desiccation tolerance. Treated seeds are usually redried before use, but they would exhibit rapid germination when re‐imbibed under normal or stress conditions. Various seed priming techniques have been developed, including hydropriming (soaking in water), halopriming (soaking in inorganic salt solutions), osmopriming (soaking in solutions of different organic osmotica), thermopriming (treatment of seed with low or high temperatures), solid matrix priming (treatment of seed with solid matrices), and biopriming (hydration using biological compounds). Each treatment has advantages and disadvantages and may have varying effects depending upon plant species, stage of plant development, concentration/dose of priming agent, and incubation period. In this article, we review, evaluate, and compare effects of various methods of seed priming in improving germination of different plant species under saline and non‐saline conditions. We also discuss the known metabolic and ultra‐structural changes that occur during seed priming and subsequent germination. To maximize the utility of various seed priming techniques, factors affecting their efficiency must be examined and potential benefits and drawbacks determined. For example, quality of the seed before treatment, concentration/dose of priming agent, time period for priming, and storage quality of the seed following priming treatment must be carefully determined. Furthermore, such assessments must be based on large‐scale experiments if seed priming is to be used for large‐scale field planting. A better understanding of the metabolic events that take place in the seed during priming and subsequent germination will improve the effective application of this technology. The incorporation of advanced molecular biology techniques in seed research is vital to the understanding and integration of multiple metabolic processes that can lead to enhanced seed germination, and consequently to improved stand establishment and crop yield under saline and non‐saline conditions.

Pre‐Sowing Seed Treatment—A Shotgun Approach to Improve Germination, Plant Growth, and Crop Yield Under Saline and Non‐Saline Conditions

Tomato and salinity

Seed priming is a pre-sowing treatment which leads to a physiological state that enables seed to germinate more efficiently. The majority of seed treatments are based on seed imbibition allowing the seeds to go through the first reversible stage of germination but do not allow radical protrusion through the seed coat. Seeds keeping their desiccation tolerance are then dehydrated and can be stored until final sowing. During subse‐ quent germination, primed seeds exhibit a faster and more synchronized germination and young seedlings are often more vigorous and resistant to abiotic stresses than seedlings obtained from unprimed seeds. Priming often involves soaking seed in predetermined amounts of water or limitation of the imbibition time. The imbibition rate could be somehow controlled by osmotic agents such as PEG and referred as osmopriming. Halopriming implies the use of specific salts while "hormopriming" relies on the use of plant growth regulators. Some physical treatments (UV, cold or heat,..) also provide germination improvement thus suggesting that priming effects are not necessarily related to seed imbibition. A better understanding of the metabolic events taking place during the priming treatment and the subsequent germination should help to use this simple and cheap technology in a more efficient way.

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Seed Priming: New Comprehensive Approaches for an Old Empirical Technique

RAPDs (Randomly Amplified Polymorphic DNAs) were used to discriminate among 23 cultivars of oilseed rape (Brassica napus) selected from several breeding programs. A set of 100 random sequence 10-mer primers were tested, of which 70 produced bands and 22 showed evidence of polymorphism. A selection of six primers produced 23 polymorphic bands of between 300 to 2200 base pairs in size, sufficient to distinguish between the cultivars. An analysis of seed of five cultivars obtained from four different sites showed stability of banding pattern over source of seed. The analysis was repeated using four different thermocyclers, each of which produced the same band pattern. UPGMA cluster analysis indicates that the relationships among some of the cultivars is closer for those from the same breeding program than for those from different programs. The results of this study show that RAPDs can be used as a method of identification for oilseed rape cultivars.

Discrimination among cultivars of rapeseed (Brassica napus L.) using DNA polymorphisms amplified from arbitrary primers

Review on dormancy, germinability and germination in crop and weed seeds.

Studies based on X-ray photographs were conducted to predict the morphology of tomato ( Lycopersicon esculentum Mill.) seedlings at transplanting stage. Currently, seed-lot quality of tomato seeds for growing commercial transplants is determined with grow-out tests in the greenhouse because the standard germination test fails to predict the percentage of normal or usable transplants (UTs). These grow-out tests, however, are difficult to standardize. An X-ray evaluation procedure is presented as an alternative. X-ray images nondestructively provide information on embryo size and morphology and the amount of endosperm and the area of free space. These parameters correlate well with the morphology of 14-day old seedlings. Cotyledon morphology has the highest correlation with the percentage of UTs. A test based on the evaluation of X-ray images, classifying the cotyledon morphology and seed free space, predicts the percentage of UTs more accurately than the currently used germination test. A second method based on an equation that uses the probabilities of all X-ray categories proportionally predicts the percentage of UTs of primed seeds more accurately than the first method. Selecting individual seeds based on X-ray images has the potential to raise the percentage of UTs of seed lots. On the average, the percentage of UTs of control seeds was 22% higher after hand selection based on X-ray evaluation. Primed seeds gave 12% higher results. Hence, X-ray analysis can predict seedling performance and enable the selection of high-quality seeds.

/pdf/predicting-tomato-seedling-morphology-by-x-ray-analysis-of-2pnpge9ebe.pdf

Predicting Tomato Seedling Morphology by X-ray Analysis of Seeds

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.

X-ray studies on changes in embryo and endosperm morphology during priming and imbibition of tomato seeds

Rapid identification techniques used in laboratories of the International Seed Testing Association : a survey

Non-destructive testing is important in the search for seed characteristics that relate to quality. It provides a means for consecutive testing on a seed by seed basis. If the tests are fast and can be automated they can form the basis for rapid new analysis methods or online sorting. X-ray analysis of tomato seeds enables us to predict seedling morphology. Acoustic analysis provides rapid physical assessment of seeds, which correlates with seed coat conditions. Both techniques can be automated and provide potential new sorting techniques. The X-ray technique provided us with some evidence for the origin of vigour differences in tomato seeds: morphological differences in seed embryos and the availability of endosperm proved to be crucial.

R. A. van Zwol

Papers

Predicting Tomato Seedling Morphology by X-ray Analysis of Seeds

X-ray studies on changes in embryo and endosperm morphology during priming and imbibition of tomato seeds

Rapid identification techniques used in laboratories of the International Seed Testing Association : a survey

Non-destructive seed evaluation with impact measurements and X-ray analysis

Purity analysis of Festuca rubra and F. ovina seed samples by densitometric evaluation of thin-layer chromatographic patterns