Seed dormancy is an innate seed property that defines the environmental conditions in which the seed is able to germinate. It is determined by genetics with a substantial environmental influence which is mediated, at least in part, by the plant hormones abscisic acid and gibberellins. Not only is the dormancy status influenced by the seed maturation environment, it is also continuously changing with time following shedding in a manner determined by the ambient environment. As dormancy is present throughout the higher plants in all major climatic regions, adaptation has resulted in divergent responses to the environment. Through this adaptation, germination is timed to avoid unfavourable weather for subsequent plant establishment and reproductive growth. In this review, we present an integrated view of the evolution, molecular genetics, physiology, biochemistry, ecology and modelling of seed dormancy mechanisms and their control of germination. We argue that adaptation has taken place on a theme rather than via fundamentally different paths and identify similarities underlying the extensive diversity in the dormancy response to the environment that controls germination.

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Seed dormancy and the control of germination

The endosperm is a barrier for radicle protrusion of many angiosperm seeds. Rupture of the testa (seed coat) and rupture of the endosperm are two sequential events during the germination of Lepidium sativum L. and Arabidopsis thaliana (L.) Heyhn. Abscisic acid (ABA) specifically inhibits the endosperm rupture of these two closely related Brassicaceae species. Lepidium seeds are large enough to allow the direct measurement of endosperm weakening by the puncture force method. We found that the endosperm weakens prior to endosperm rupture and that ABA delays the onset and decreases the rate of this weakening process in a dose-dependent manner. An early embryo signal is required and sufficient to induce endosperm weakening, which afterwards appears to be an organ-autonomous process. Gibberellins can replace this embryo signal; de novo gibberellin biosynthesis occurs in the endosperm and weakening is regulated by the gibberellin/ABA ratio. Our results suggest that the control of radicle protrusion during the germination of Brassicaceae seeds is mediated, at least in part, by endosperm weakening. We propose that Lepidium is an emerging Brassicaceae model system for endosperm weakening and that the complementary advantages of Lepidium and Arabidopsis can be used in parallel experiments to investigate the molecular mechanisms of endosperm weakening.

/pdf/endosperm-limited-brassicaceae-seed-germination-abscisic-466prjlylm.pdf

Endosperm-limited Brassicaceae Seed Germination: Abscisic Acid Inhibits Embryo-induced Endosperm Weakening of Lepidium sativum (cress) and Endosperm Rupture of Cress and Arabidopsis thaliana

Preplant Physiological Seed Conditioning

Secondary lichen products can be extracted from air-dry thalli of Xanthoria parietina, Xanthoparmelia conspersa and Parmelina tiliacea with 100% acetone without affecting either short-or long-term viability. In Xanthoria parientina damage by acetone started to occur as water content reached the critical lower limit for photosystem II (PSII) activity. Extraction of the blue-light absorbing cortical pigment parietin increased the susceptibility of both air-dry and hydrated thalli to high light. Damage by high light levels caused a permanent reduction in Fv/Fm, quantum yield for photosynthetic O2 production and photosynthetic capacity measured after a 2-day recovery period at low light levels (20 μmol photons m-2 s-1). Parietin therefore protects the photosynthetic apparatus of Xanthoria parietina against damage by high light levels. Extraction of UV-absorbing pigments from Xanthoparmelia conspersa and Parmelina tiliacea did not increase photoinhibition after 24 h exposure to high light.

Parietin, a photoprotective secondary product of the lichen Xanthoria parietina.

Presowing Seed Priming

Osmotic conditioning of seeds: physiological and biochemical changes

Acetone is able to carry chemicals through the seed coats to reach the embryo. The amount of chemical penetrating into the seeds depends on the seed type, permeation time, and the concentration of solution. The chemicals reaching the embryo are biologically effective.

Penetration of dry seeds with chemicals applied in acetone.

The forces required to puncture intact lettuce (Lactuca sativa) seed and pericarp, endosperm and embryo were measured by the Instron Universal Testing Machine. It required about 0.6 newton to puncture the endosperm in seeds imbibed in the dark at 6, 12 and 24 hours. Endosperm of seeds imbibed in the light or in dark with gibberellic acid required about 4.2 newtons at 6 and at 12 hours and only about 0.15 newton at 24 hours. Forces required to puncture embryo at all treatments and times remained constant at about 0.3 newton. Changes in the strength of the endosperm do not appear to be related directly to protrusion of the radicle.

Changes in the Strength of Lettuce Endosperm during Germination

Intact lettuce seed germination was inhibited by cordycepin but not by actinomycin D; however, when seeds were clipped at the cotyledonary end, actinomycin D partially inhibited germination. Uptake studies with intact seeds using 3H-actinomycin D showed that it was unable to reach the embryo prior to radical protrusion. 3H-Cordycepin uptake studies using intact seeds showed that cordycepin was able to reach the embryo during the first 3 hours of incubation and at subsequent times. The pericarp and endosperm offered resistance to penetration of cordycepin into the embryo. In contrast to actinomycin D, cordycepin markedly inhibited 3H-uridine incorporation into RNA of intact seeds during the first 10 and 12 hours of incubation. About 60% of 3H-adenosine incorporation into poly A-RNA was inhibited by cordycepin during 12 hours of incubation, whereas actinomycin D had little effect. RNA synthesis appears to be essential for seed germination.

Differential Effects of Actinomycin D and Cordycepin in Lettuce Seed Germination and RNA Synthesis

The number of isoperoxidases and the intensity of certain isozymes increased with increasing periods of stratification of pear (Pyrus communis cv. Bartlett) embryos. The presence of GA3 or kinetin during stratification enhanced the activity of certain isoperoxidases, and these enhancements were blocked in the presence of ABA which by itself had an inhibitory effect. Enhancement in isoperoxidases of pear embryos during stratification was inhibited by 6-methylpurine and cycloheximide; and in the presence of either of these two inhibitors, stratification failed to release the dormancy. Pear embryos germinated for 3 days showed changes in the pattern of isoperoxidases depending on the length of stratification.

/pdf/changes-in-isoperoxidases-during-cold-treatment-of-dormant-4nr28nlp6l.pdf

Kar-Ling Tao

Papers

Osmotic conditioning of seeds: physiological and biochemical changes

Penetration of dry seeds with chemicals applied in acetone.

Changes in the Strength of Lettuce Endosperm during Germination

Differential Effects of Actinomycin D and Cordycepin in Lettuce Seed Germination and RNA Synthesis

Changes in Isoperoxidases during Cold Treatment of Dormant Pear Embryo