Current interest in seed hydration treatments for the improvement of level, rate and uniformity of germination or field emergence has revealed how little is known of the physiology of germination control under water stress. This review surveys briefly the responses of seeds held at different hydration levels, from normal germination in a free water supply to seed activation without germination under slight moisture stress, seed deterioration at greater moisture stress and the damage that can be caused to seeds in very dry conditions, as well as the responses to subsequent dehydration. Inhibition of germination, though not of seed activation, at certain levels of water stress is likened to various forms of dormancy, and the mechanism governing the initiation of cell elongation is suggested as the possible key to control over germination. Several lines of evidence on cell membrane integrity and action, and their responses to external factors, point to the role that the membrane may play in cell elongation (and hence germination) control, and membrane integrity may also be associated with the transition between seed deterioration at one hydration level and seed activation and repair at slightly higher hydration levels. Seed activation without germination is also considered in an ecological context.

The physiology of seed hydration and dehydration, and the relation between water stress and the control of germination: a review

Seed Research-Past,Present and Future.- Use of Transgenic Plants for Studies of Seed-Specific Gene Expression.- Impact of Simulated Drought Stress on Protein Body Cycle in Radicles of Developing and Germinating Cotton Seeds.- Synthetic Seed for Clonal Production of Crop Plants.- Synthesis of Acid-Soluble,ABA-Inducible Proteins in Wheat Embryos from Dormant Grain.- Hormones, Genetic Mutants and Seed Development.- Membrane Behavior in Drought and Its Physiological Significance.- The Basis of Recalcitrant Seed Behaviour: Cell Biology of the Homoiohydrous Seed Condition.- Phytin Synthesis and Deposition.- Calcium-Regulated Metabolism in Seed Germination.- Transport and Targeting of Proteins to Protein Storage Vacuoles (Protein Bodies) in Developing Seeds.- Biochemical Adaptations to Anoxia in Rice and Echinochloa Seeds.- Some Aspects of Metabolic Regulation of Seed Germination and Dormancy.- Phytochrome and Sensitization in Germination Control.- The Role of Light and Nitrate in Seed Germination.- Water-Impermeable Seed Coverings as Barriers to Germination.- Control of Germination and Early Development in Parasitic Angiosperms.- Factors Eliciting the Germination of Photoblastic Kalanchoe Seeds.- Factors Influencing the Efficacy of Dormancy-Breaking Chemicals.- Physiological Mechanisms Involved in Seed Priming.- Germination Research Towards the Nineties: A Summary and Prognosis.

Recent advances in the development and germination of seeds

A brief survey of the biochemistry of the alternative oxidative pathway (‘cyanide-resistant respiration’) and its occurrence in vivo is given. Several hypotheses about the physiological significance of the alternative chain are discussed. These include a role in (1) heat production, (2) fruit ripening, (3) respiration of plants that contain high levels of cyanogenic glycosides, producing HCN upon wounding, (4) oxidation of NADH that is produced by various causes in excess of that required for ATP production, (5) ion uptake, and (6) osmoregulation.

In intact roots of higher plants, the activity of the alternative pathway is more active when less carbohydrate is required for assimilation of N (NH+4 NO-3 or N2) and is less active in those when carbohydrates are being stored in a storage organ or when the availability of photosynthate is reduced. An increase in carbohydrate requirement for osmoregulation is also correlated with decreased alternative chain activity.

It is concluded that the alternative pathway in roots plays an important role in oxidation of sugars which are not required for carbon skeletons, energy production for growth and maintenance processes, osmoregulation or storage. However, the significance of this role may vary in different tissues and physiological states.

The physiological significance of cyanide‐resistant respiration in higher plants

The cereal aleurone layer is a secretory tissue whose function is regulated by gibberellic acid (GA) and abscisic acid (ABA). Aleurone cells lack functional chloroplasts, thus excluding photosynthesis as a source of active oxygen species (AOS) in cell death. Incubation of barley aleurone layers or protoplasts in GA initiated the cell death programme, but incubation in ABA delays programmed cell death (PCD). Light, especially blue and UV-A light, and H(2)O(2) accelerate PCD of GA-treated aleurone cells, but ABA-treated aleurone cells are refractory to light and H(2)O(2) and are not killed. It was shown that light elevated intracellular H(2)O(2), and that the rise in H(2)O(2) was greater in GA-treated cells compared to cells in ABA. Experiments with antioxidants show that PCD in aleurone is probably regulated by AOS. The sensitivity of GA-treated aleurone to light and H(2)O(2) is a result of lowered amounts of enzymes that metabolize AOS. mRNAs encoding catalase, ascorbate peroxidase and superoxide dismutase are all reduced during 6-18 h of incubation in GA, but these mRNAs were present in higher amounts in cells incubated in ABA. The amounts of protein and enzyme activities encoded by these mRNAs were also dramatically reduced in GA-treated cells. Aleurone cells store and metabolize neutral lipids via the glyoxylate cycle in response to GA, and glyoxysomes are one potential source of AOS in the GA-treated cells. Mitochondria are another potential source of AOS in GA-treated cells. AOS generated by these organelles bring about membrane rupture and cell death.

Active oxygen and cell death in cereal aleurone cells.

Oxygen uptake during the first hours of imbibition in intact soybean and mung bean seeds showed a marked sensitivity to potassium cyanide but was unaffected by addition of either salicylhydroxamic acid or propyl gallate. However O(2) uptake by finely ground seed particles was very sensitive to the addition of either compound. The results indicated that O(2) uptake in intact, imbibing seeds was associated with a cyanide-sensitive process, most probably mitochondrial mediated respiration, and not the result of the cyanide-insensitive lipoxygenase activity which was readily detectable in ground seed particles.The antioxidant propyl gallate was found to inhibit specifically alternative pathway electron transfer in isolated mung bean mitochondria. Half-maximal inhibition occurred with 2 to 5 micromolar propyl gallate. Kinetic analysis indicated that propyl gallate inhibition of the alternative pathway occurred at, or very near, the site of inhibition of the alternative pathway by salicylhydroxamic acid.A high level of lipoxygenase activity was found to be associated with washed mitochondria isolated from a variety of etiolated plant tissues. Most of this lipoxygenase activity could be eliminated from mung bean mitochondria if the mitochondria were purified on a discontinuous sucrose gradient. This indicated that the mitochondrial-associated activity was probably the result of nonspecific adsorption of lipoxygenase onto the mitochondrial membranes during isolation.

/pdf/alternative-respiratory-pathway-its-role-in-seed-respiration-4ecrqimk0q.pdf

Alternative Respiratory Pathway: ITS ROLE IN SEED RESPIRATION AND ITS INHIBITION BY PROPYL GALLATE.

Experiments with germinating seeds of Wayne soybean (Glycine max Merr.) show that between the 4th and the 8th hour of germination, respiration experiences a transition from predominantly "alternate" respiration, which is sensitive to salicylhydroxamic acid, to a cyanide-sensitive respiration. The dependence of early germination stages on alternate respiration is reflected in several types of seed functions, including subsequent root growth rate, chlorophyll synthesis, and germination itself. The early period of germination is shown to require a normal O(2) tension, which is no longer a requirement at later stages. The changing sensitivity to cyanide and to salicylhydroxamic acid is found to be common to seven different types of germinating seeds. It is proposed that the alternate pathway of respiration provides something essential for the completion of the earliest stages of seed germination.

/pdf/respiratory-transition-during-seed-germination-dacin8mw79.pdf

Respiratory Transition during Seed Germination.

Phytochrome-mediated germination of Lactuca sativa L. cv. Waldmann's Green seeds was inhibited strongly by 10 millimolar salicylhydroxamic acid (SHAM), but only slightly delayed by the same level of KCN. SHAM was most effective if applied within the 8-hour potentiation period (release from dormancy) following red light treatment, but much less effective with completely potentiated seeds. SHAM at 3 millimolar actually hastened completion of potentiation, whereas concentrations of 6.6 millimolar or higher retarded the process. A temporary upsurge of O(2) consumption was particularly evident during the period of most rapid potentiation (3 hours after red light), especially in the seed sections containing the embryonic axis. The embryonic axis obtained from dormant seeds also contained most of the SHAM-sensitive O(2) uptake. However, 8 hours of potentiation caused loss of SHAM sensitivity from axes and a simultaneous gain of SHAM sensitivity by cotyledons. Concomitant with this increased sensitivity to SHAM, O(2) uptake by cotyledonary tissues lost some sensitivity to KCN. Red light-stimulated metabolic processes leading to germination were blocked more effectively by SHAM than by KCN, but O(2) consumption by both dormant and nondormant seeds was much less sensitive to 10 millimolar SHAM than to the same concentration of KCN. This apparent contradiction between effects of SHAM on potentiation and O(2) uptake may be a result of: (a) compensatory electron flow through the cytochrome pathway at the expense of the alternate pathway; (b) a functional site of action of SHAM that differs from the organized, energy-coupled respiratory system; or (c) a combination of these possibilities.

Semahat Yentur

Papers

Respiratory Transition during Seed Germination.

Cyanide-insensitive, Salicylhydroxamic Acid-sensitive Processes in Potentiation of Light-requiring Lettuce Seeds