Salt tolerance of barley (Hordeum vulgare L.) cultivars at the germination stage: Analysis of the response functions

Two models, initially proposed by Van Genuchten (1983) for evaluating salinity-yield response curves at the adult stage, were applied to study the salinity response of 24 barley cultivars at the germination stage. According to the calculated salinity threshold, EC_t (the solution electrical conductivity, EC, at which germination starts to decrease), and EC₅₀ (the solution EC at which germination is reduced by 50%) parameters, both models give similar results, although model 2, a sigmoid-form curve, fits the observed data slightly better than model 1, a piecewise response function. Also, the results suggest that, for model 1, EC_t seems to be the most reliable parameter for screening barley germplasm because it clearly discriminates the relative salt-tolerance of the studied cultivars and, furthermore, it basically determines their salinity response for the 100 to 50% germination interval. On the other hand, the model 1 s parameter — percent germination decrease per unit salinity increase bove EC_t—is less relevant because of its smaller variation interval and lack of correlation with EC₅₀, indicating that the salinity response of the studied cultivars for the 50% germination value is independent of this parameter.     

Osmotic sensitivity in relation to salt sensitivity in germinating barley seeds

Abstract Cultivars of barley (Hordeum vulgare L.) were tested for germination sensitivity to progressively higher concentrations of salt, mannitol, and betaine. The three solutes were equally inhibitory at equal osmotic potential, but there was a consistent difference in osmotic sensitivity between two cultivars, CM-67 and Briggs (Briggs was the most sensitive). There was no difference between the two cultivars in salt or water uptake from salt solutions during imbibition. Brief presoaking in water did not improve salt resistance, indicating that a hydration-dependent decrease in membrane permeability is not involved in salt tolerance. The calcium content of Briggs was higher than CM-67. These results suggest that salt inhibits barley germination primarily by osmotic effects, and that salt influx during imbibition does not play a role in this inhibition. A hypothesis regarding salt effects on germination is discussed.     

Effect of environmental factors on seed germination and emergence of Lepidium vesicarium

Lepidium vesicarium is a weed species with a wide distribution in the rangelands and dry‐land farming in East Azarbaijan, Iran. The experiments were undertaken to assay the effects of light, temperature, pH, osmotic potential, NaCl concentration and burial depth on seed germination and emergence of L. vesicarium. Germination was maintained at high levels (> 80%) over a wide day/night temperature range (10/5 to 30/20°C), but a severe reduction in the germination rate of L. vesicarium was found below 20/10°C. Germination of L. vesicarium was influenced by different light/dark regimes, as the germination rate was highest at 16 h light for the all treatments (0, 8, 12, 16 and 24 h light). Germination was 92–95% over a wide range of pH (2‐10). Germination was >50% at a water potential of ?0.7 MPa and salinity of 21 dS/m, indicating that drought and salt conditions have a minimal impact on seed germination. With increasing burial depth from 0 to 2 cm, the number of days required for 50% emergence increased and no germination was observed at burial depths deeper than 3 cm. This suggests that L. vesicarium would become troublesome in the rangelands and for growers in reduced‐tillage cropping systems. The ability to emerge from shallow depths, coupled with tolerance of a wide pH range, drought and salinity at germination, should be taken into account when managing this weed species.     

The influence of aggregate size and drannage potential on germination of barley seeds in flooded soil

Emergence and growth of barley was severely decreased by short periods (less than 24 hours) of pre-emergence waterlogging at 20°C. The extent of damage depended on a combination of duration of waterlogging, soil water potential and aggregate size. Potentials of less than—4kPa prevented loss of plants developing in aggregates of less than 2 mm diameter after a transitory period of waterlogging although some shoot and root damage occurred. By comparison seeds growing in soil consisting of aggregates greater than 2 mm in diameter were not damaged by transitory waterlogging even when drainage only occurred at−0.8kPa. The severity of damage increased with the period of waterlogging. A criterion obtained as the product of mean size grade and water potential gave a single value (−4NM⁻¹) below which emergence was satisfactory. Waterlogging halfway through germination gave more severe damage than near sowing date or near emergence.     

Effects of NaCl salinity in vivo and in vitro on ribonuclease activity of Vigna unguiculata cotyledons during germination

Pitiuba bean [ Vigna unguiculata (L.) Walp.] seeds were sown in water or. in 0.1 M NaCl. Seedling growth and cotyledon nucleic acid mobilization were delayed by NaCl salinity. The differences in cotyledonary RNase activity between seeds sown in water and in NaCl solutions suggest that salinity delays the activation and/or de novo synthesis of the enzyme. Cotyledon extracts were subjected to gel filtration through Sephadex G-100, and RNase activity measured. Only one cotyledonary RNase appeared during germination, and salinity did not induce any change in molecular weight of the enzyme. Salinity inhibited 45% of the specific activity of the RNase on the 5th day of the experimental period. The same salt concentration (0.1 M NaCl) added in vitro inhibited only 8 % of the specific activity of the enzyme. This difference may indicate that NaCl in vivo affects mainly the de novo synthesis of the RNase.     

Osmotic and specific ion effects on the germination of Prosopis strombulifera

BACKGROUND AND AIMS: Salinity can affect germination of seeds either by creating osmotic potentials that prevent water uptake or by toxic effects of specific ions. Most studies have only used monosaline solutions, although these limit the extent to which one can interpret the results or relate them to field conditions. The aim of this work was to evaluate the germination of Prosopis strombulifera seeds under increasing salinity by using the most abundant salts in central Argentina in monosaline or bisaline iso-osmotic solutions, or in solutions of mannitol and polyethylene glycol. METHODS: Seeds were allowed to germinate under controlled conditions in a germination chamber at 30 +/- 1 degrees C and at 80 % r.h. Salinizing agents were KCl, NaCl, Na(2)SO(4), K(2)SO(4), NaCl + Na(2)SO(4) and KCl + K(2)SO(4) and osmotic agents were polyethylene glycol 6000 and mannitol. Treatments for all osmotica consisted of 0.0, -0.4, -0.8, -1.2, -1.5, -1.9 and -2.2 MPa solutions. KEY RESULTS: The percentage of germination decreased as salinity increased. SO(4)(2-) in monosaline solutions, with osmotic potentials -1.2 MPa and lower, was more inhibitory than Cl(-) at iso-osmotic concentrations. This SO(4)(2-) toxicity was alleviated in salt mixtures and was more noticeable in higher concentrations. K(+) was more inhibitory than Na(+) independently of the accompanying anion. CONCLUSIONS: Different responses to different compositions of iso-osmotic salt solutions and to both osmotic agents indicate specific ionic effects. This study demonstrates that the germination of P. strombulifera is strongly influenced by the nature of the ions in the salt solutions and their interactions. Comparative studies of Cl(-) and SO(4)(2-) effects and the interaction between SO(4)(2-) and Cl(-) in salt mixtures indicate that extrapolation of results obtained with monosaline solutions in the laboratory to field conditions can be speculative.     

Allelopathic effect of Cistus ladanifer on seed germination

1. The allelopathic effect of the exudate secreted by Cistus ladanifer leaves was tested on different plant species. Cynodon dactylon and Rumex crispus , species absent from C. ladanifer scrub (jarales), were clearly inhibited by the exudate.
2. Species present in the jarales, Medicago polymorpha and Lolium rigidum , showed no direct inhibition of germination, but the process this was delayed, and cotyledon and root size was diminished. This implies inhibition of seedling growth, and may explain the low abundance of these species inside the jarales.
3. The active constituents in the inhibition of germination are low molecular weight phenolic compounds. The flavonoids appear to exert no direct effect on germination of the species tested, but do provoke a reduction in cotyledon and root size, and could thereby inhibit subsequent seedling development.     

The effect of phenols on respiratory enzymes in seed germination

Low molecular weight phenolic compounds were identified in two soilswith different vegetative cover, Fagus sylvatica, L. andPinus laricio, Poiret, spp. calabrica, and were tested atdifferent concentrations on seed germination of Pinuslaricio, and on respiratory and oxidative pentose phosphate pathwayenzymes involved in the first steps of seed germination. The data obtained showthat there are marked differences in the phenolic acid composition of the twoinvestigated soils. All the phenolic compounds bioassayed inhibited seedgermination and those extracted from Pinus laricio soilwere particularly inhibitory. We also found that the non-germination of seedsisstrongly correlated to the inhibition of the activities of enzymes ofglycolysisand the oxidative pentose phosphate pathway.     

Reversing the inhibitory effect of paclobutrazol on seed germination of Amaranthus paniculatus by GA3, ethephon or ACC

The germination of Amaranthus paniculatus seeds was inhibited by applying paclobutrazol, a specific inhibitor of gibberellin biosynthesis. This inhibition was markedly counteracted by gibberellin A₃ (GA₃), suggesting that endogenous gibberellins are required for germination in this species. The inhibitory effect of paclobutrazol was also overcome by ethephon (2-chloroethylphosphonic acid) or the precursor of ethylene biosynthesis, ACC (1-aminocyclopropane-l-carboxylic acid). Thus the physiological effect of gibberellin can be mimicked by ethylene released from ethephon or synthesised from exogenous ACC. It is suggested, that endogenous gibberellins are involved in germination of Amaranthus paniculatus seeds and that action of GA₃ can be substituted by ethylene.Abbreviations ACC 1-aminocyclopropane-l-carboxylic acid - AMO-1618 (2-isopropyl-5methyl-4-trimethylammoniumchloride)-phenyl-l-piperidinium-carboxylate - ancymidol -cyclopropyl--(4-methoxyphenyl)-5-pyrimidine methanol - chloromequat chloride (2-chloroethyl)trimethylammoniumchloride - ethephon 2-chloroethylphosphonic acid - GA gibberellin A₃ - paclobutrazol (2RS, 3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-lyl)pentan-3-ol - Phosphon D 2,4,dichlorobenzyl-tributhylphosphoniumchloride - tetcyclacis 5,(4-chlorophenyl)-3,4,5,9,10-pentaaza-tetracyclo)5,4,1,0,Z,6,0^8,11 dodeca-3,9-diene     

Differential effects of four abiotic factors on the germination of salt marsh annuals

Interspecific differences in responsiveness to temperature, photoperiod, soil salinity, and soil moisture confirm the hypothesis that abiotic factors differentially affect the germination of salt marsh plants. In growth chamber experiments, four of eight annual species responded to small differences in temperature or photoperiod. Increasing soil salinity decreased the final proportion of seeds germinating and slowed germination for each of the seven species tested. Higher soil moisture increased the proportion germinating of five species and germination speed of all seven species. Salinity and moisture interacted to affect the proportion germinating of five species and germination speed of all seven species. Although the abiotic factor with the largest effect on germination varied among species, more species responded to, and the magnitudes of the responses were larger for, soil salinity than for the other abiotic factors. These germination tests partially explained interspecific differences in the timing of germination in the field. Patterns of Hutchinsia procumbens, Lythrum hyssopifolium, Parapholis incurva, and possibly Lasthenia glabrata ssp. coulteri germination in response to a nonseasonal rainfall could be explained by their response to salinity, temperature, or photoperiod. Fine-scale differences in the timing of establishment within the typical germination window and spatial distributions along salinity and moisture gradients were not explained.     

Salinity and salt composition effects on seed germination and root length of four sugar beet cultivars

Salinization is one of the most important factors affecting agricultural land in the world. Salinization occurs naturally in arid and semiarid regions where evaporation is higher than rainfall. Sugar beet yield declines with an increase in salinity, but the sensitivity to salts varies with salt composition in water and sugar beet growth stage. The aim of this study was to determine the effect of water salinity levels and salt composition on germination and seedling root length of four sugar beet cultivars (PP22, IC2, PP36, and 7233). The experiments were undertaken with irrigation water with two salt compositions (NaCl alone and mixture of MgSO₄ + NaCl + Na₂SO₄ + CaCl₂) in three replicates. Thirteen salinity levels with electrical conductivity (EC) of the irrigation water ranging from 0 to 30 dS/m were applied to each cultivar in both experiments. Seed germination percentage and seedling root length growth were determined in 13 days. Statistical analysis revealed that germination and root length were significantly affected by salt composition, cultivars and salinity levels. Regardless of salt composition, seed germination and seedling root length were significantly affected by the irrigation water with EC up to 8 dS/m and 4 dS/m, respectively. Except for cultivar PP22, the adverse effect of salinity of the irrigation water on seed germination and seedling root length was higher for NaCl alone than for the salt mixture, which refers to lower salt stress in field conditions with natural salt composition. Presented at the International Conference on Bioclimatology and Natural Hazards, Poľana nad Detvou, Slovakia, 17–20 September 2007.     

Location of sugars in barley seeds during germination by NMR microscopy

The distribution and fluctuation of sugars in germinating barley seeds were examined by ¹³C nuclear magnetic resonance (NMR) spectroscopy, ¹H-NMR imaging and ¹H-NMR localized spectroscopy in relation to morphology. Maltose, sucrose, fructose and oils were detected in intact imbibed seeds by ¹³C-NMR spectra. During the first 6 d of germination, the maltose content increased and the oil content gradually decreased, whilst the levels of sucrose and fructose remained constant. Sugars were located by ¹H-NMR images and ¹H-NMR localized spectra in the vascular bundle of the seeds as well as in the solubilized endosperm. They were also detected in the shoots. The sugars detected in an 80% ethanol shoot extract were sucrose and glucose, which were located in the vascular bundles but not in the mesophyll cells of the coleoptile. They were also located in the basal part of the shoot, but not above 7 mm from the scutellum. The data suggest that the sugars are primarily transported through the vascular bundles and, at the same time, rapidly incorporated into mesophyll cells in the leaves.     

Soil salinity delays germination and limits growth of hyphae from propagules of arbuscular mycorrhizal fungi

Colonisation of plant roots by some arbuscular mycorrhizal (AM) fungi is reduced in the presence of sodium chloride (NaCl), probably due to a direct effect of NaCl on the fungi. However, there appear to be differences between the fungi in their ability to colonise plants in the presence of NaCl. This experiment tested the hypothesis that propagules of different isolates and species of AM fungi from saline and nonsaline soils would differ in their ability to germinate and grow in the presence of NaCl in the soil solution. Spores or pieces of root colonised by a range of AM fungi were incubated between filters buried in soil to which NaCl had been added at concentrations of 0, 150 or 300 mM in the soil solution. At regular intervals, filters were removed from the soil and both the percentage of propagules which had germinated and the length of proliferating hyphae were determined. Germination of spores of AM fungi studied was delayed in the presence of NaCl, but the fungi differed in the extent to which germination was inhibited. Two isolates of Scutellospora calospora reached maximum germination in 300 mM NaCl, but neither of two isolates of Acaulospora laevis germinated in the presence of NaCl. Germination of spores of the other fungi, including some isolated from saline soil, fell between these extremes. For some fungi, the specific rate of hyphal extension was reduced by NaCl. For others, the specific rate of growth was similar in the presence of NaCl to that in the control treatment, but overall production of hyphae was reduced in the NaCl treatments because germination was reduced.     

The in vitro effect of gossypol and its interaction with salts on conidial germination and viability of Fusarium oxysporum sp. vasinfectum isolates

AIMS: To assess the effect of different concentrations of gossypol (0, 2, 4, 10 and 20 mg l(-1)) in combination with NaCl and Na(2)SO(4) (20 mS cm(-1)) on the conidial germination and viability of Fusarium oxysporum f.sp. vasinfectum (Fov). METHODS AND RESULTS: A multinomial logistic model was developed to estimate the germination probability of Fov. The inhibitory effect was markedly evident at the two highest concentrations of gossypol; it varied among the isolates tested and with time, and it was attenuated by the presence of sodium salts. The inhibition was temporary as the germination probability increased after 8 h. Fluorescent staining revealed that gossypol either killed the conidia or retarded the elongation of the germ tubes. CONCLUSION: Fov showed the ability to overcome gossypol inhibition over time, and the inhibitory effect is reduced under saline conditions. Differential responses among Fov isolates to the presence of gossypol suggest that gossypol tolerance is genetically determined in the pathogen. SIGNIFICANCE AND IMPACT OF THE STUDY: This study suggests that selecting for high plant gossypol cultivars would have minimal effect on the overall Fov resistance of cotton. A new statistical model was developed to explore the statistical significance of plant-pathogen interactions.     

Effects of salinity and temperature on ex situ germination of the threatened Gulf of St. Lawrence Aster,Symphyotrichum laurentianum Fernald (Nesom)

The threatened Gulf of St. Lawrence Aster, Symphyotrichum laurentianum Fernald (Nesom), is an annual coastal halophyte of the southern Gulf of St. Lawrence, Canada. We examined the effects of salinity (0–20 g/L) and temperature (16–30°C) on germination of S. laurentianum seeds over 32 days. The time‐course of germination was significantly affected by both salinity and temperature. At lower temperatures (16°C and 23°C), germination was inhibited by salt water at days 16 and 32. However, at 30°C germination rates after 16 days were highest at an intermediate salinity, whereas after 32 days germination was uniformly high in all salinity treatments. Overall, the effect of temperature on germination was much stronger than the effect of salinity. Delays in germination resulting from exposure to salinity or from low soil temperatures could set up strong size asymmetries between seedlings of S. laurentianum and the surrounding vegetation, leading to suppression of growing seedlings via shading. Because germination has the potential to be a significant population bottleneck for this seed‐dependent annual, conservation efforts should consider microsite suitability for germination in the management of natural populations and in the selection of sites for explants.     

Effects of NaCl stress on germination, antioxidant responses, and proline content in two rice cultivars

We investigated the physiological and biochemical bases for salt tolerance in two rice (Oryza sativa L.) cultivars — relatively salt-tolerant ‘Dongjin’ and salt-sensitive ‘Kumnam’. Salinized hydroponic cultures were studied at the germination and seedling stages. NaCI inhibited germination more severely in ‘Kumnam’ than in ‘Dongjin’. Increasing the salt concentration also deterred growth to a larger extent in the former. Moreover, the leaves of ‘Kumnam’ exhibited greater increases in lipid peroxidation and Na⁺ accumulation than those of ‘Dongjin’ under stress. The activities of constitutive and salt-induced superoxide dismutase (SOD, EC 1.15.1.1) and ascorbate peroxidase (AP, EC 1.11.1.11) were also higher in ‘Kumnam’, while only catalase (CAT, EC 1.11.1.6) activity was slightly higher in stressed plants of ‘Dongjin’. The positive correlation between leaf proline levels and NaCI concentration was more evident in ‘Kumnam’. However, ‘Dongjin’ seeds, which had higher germinability in the presence of NaCI, also contained more proline. These results suggest that the higher salt tolerance in ‘Dongjin’ seedlings could be ascribed to their lower NaCI accumulations in the leaves. This presumably is due to reductions in the uptake or transport rates of saline ions to the shoots from the roots. Finally, we believe that the higher germination rate by ‘Dongjin’ is caused by its higher seed proline content.     

The effect of a plant-derived smoke extract on the germination of light-sensitive lettuce seed

A simple and rapid bioassay using seeds of Lactuca sativa L. Grand Rapids has been developed for the detection of germination-enhancing compound(s) in plant-derived smoke extracts. This light-sensitive species germinates within 24 h in the dark at 20 or 25°C and shows responsiveness to smoke-derived extracts over a wide concentration range. For some seed lots where the P_fr level is high and germination in the dark is unacceptably high, a brief (10 min) exposure to far-red light, one hour after the start of imbibition in the dark, is necessary to clearly demonstrate biological activity in the smoke extracts.     

Effect of NaCl,Na<Subscript>2</Subscript>SO<Subscript>4</Subscript>, and mannitol on utilization of storage starch and formation of plastids in the cotyledons and roots of alfalfa seedlings

Utilization of storage starch in the cells of cotyledon mesophyll and root meristem in the course of alfalfa (Medicago sativa L.) seed germination on the solutions of NaCl, Na₂SO₄, and mannitol at different concentrations and identical osmotic pressure was investigated using the method of transmission electron microscopy. Ultrastructural analysis showed changes in the number of starch grains and deceleration of chloroplast development depending on the osmotic component of salt influence. At low concentrations corresponding to osmotic pressure of 202.6 kPa, Na₂SO₄ did not affect the formation of the photosynthetic machinery and utilization of starch inclusions; mannitol contributed to the preservation of considerable reserve of starch without disturbing the development of chloroplasts; NaCl did not inhibit the development of the photosynthetic machinery and induced an increase in the number of starch grains presumably at the expense of newly produced starch. When the concentration of the investigated substances increased up to the values corresponding to the osmotic pressure of 607.8 kPa, NaCl did not suppress transformation of amyloplasts into chloroplasts and utilization of starch; Na₂SO₄ inhibited the development of chloroplasts and starch utilization; mannitol decelerated transformation of amyloplasts and inhibited mobilization of starch grains. The obtained results make it possible to propose a method of preliminary estimation of tolerance of dicotyledons to abiotic stresses based on the cytological analysis of utilization of starch grains and formation of photosynthetic compartments of chloroplasts in the mesophyll of cotyledons.