Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.     

Comparison of germination and seed bank dynamics of dimorphic seeds of the cold desert halophyte Suaeda corniculata subsp. mongolica

Background and Aims

Differences in dormancy and germination requirements have been documented in heteromorphic seeds of many species, but it is unknown how this difference contributes to maintenance and regeneration of populations. The primary aim of this study was to compare the seed bank dynamics, including dormancy cycling, of the two seed morphs (black and brown) of the cold desert halophyte Suaeda corniculata and, if differences were found, to determine their influence on regeneration of the species.

Method

Seeds of the two seed morphs were buried, exhumed and tested monthly for 24 months over a range of temperatures and salinities, and germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were also investigated for the two morphs.

Key Results

Black seeds had an annual dormancy/non-dormancy cycle, while brown seeds, which were non-dormant at maturity, remained non-dormant. Black seeds also exhibited an annual cycle in sensitivity of germination to salinity. Seedlings derived from black seeds emerged in July and August and those from brown seeds in May. Seedlings were recruited from 2·6 % of the black seeds and from 2·8 % of the brown seeds in the soil, and only 0·5 % and 0·4 % of the total number of black and brown seeds in the soil, respectively, gave rise to seedlings that survived to produce seeds. Salinity and water stress induced dormancy in black seeds and decreased viability of brown seeds. Brown seeds formed only a transient soil seed bank and black seeds a persistent seed bank.

Conclusions

The presence of a dormancy cycle in black but not in brown seeds of S. corniculata and differences in germination requirements of the two morphs cause them to differ in their germination dynamics. The study contributes to our limited knowledge of dormancy cycling and seed bank formation in species producing heteromorphic seeds.     

金莲花种子的休眠、萌发与活力的研究

金莲花种子存在生理休眠现象,采用4周以上冷湿处理或不同浓度GA_3处理可以打破休眠,GA_3的最适浓度为500μg/g。不同年份收获的种子活力不同,表现在种子重量、用同样浓度GA_3处理时的发芽率及种子的最适发芽温度范围不同,1989年的种子活力低,最适发芽温度25℃左右(发芽率90％),而1990年新收的种子,从15°～25℃发芽率都高达95％以上。应用GA_3有机溶剂渗入法,可打破金莲花种子休眠促进萌发,效果良好,可以推广应用。     

The growth physics and water relations of red-light-induced germination in lettuce seeds

Using lettuce (Lactuca sativa L., cv. Grand Rapids) embryos in osmotica, we have demonstrated that when the growth rates of the embryonic axes of seeds treated with red (R) or far-red (FR) light are equalized, the axes of R-treated seeds develop a 3.4-bar decrease in water potential (paper No. III).As axial growth begins, reserve protein and phytin decrease rapidly, concomitant with increases in reducing sugars, -amino nitrogen, and inorganic and esterified soluble phosphates. However, no differences between the axes of R-and FR-treated seeds are found with respect to the changes in these compounds, indicating that these changes arise as a result of growth and are not under immediate phytochrome control. Little change in the total lipid content is found in either treatment. The axes of FR-treated seeds hydrolyze endogenous sucrose at a greater rate thant those of R-treated seeds. Axes of R-treated seeds accumulate K⁺ and Na⁺ to a greater extent than those of FR-treated seeds. When potassium salts are added to the incubation medium, R induces increased K⁺ uptake by the axis and greater medium acidification by the axis. Malate and other organic acids and acidic amino acids increase at equal rates in both treatments, indicating that inorganic anions may also be taken up to balance the ionic charges. The results are compatible with the assumption that changes in the osmotic and pressure potentials of the embryonic axes of R-treated seeds are the result of a phytochrome-stimulated proton pump which, in whole dormant seeds, would initiate water-potential changes allowing the embryos to overcome the mechanical restraint of the surrounding seed layers, resulting in germination.Abbreviations FR far-red light - PEG polyethylene glyeol 4000 - Pfr far-red-absorbing form of phytochrome - R red light III=Carpita et al. 1979     

弥勒苣苔种子的休眠萌发特性

弥勒苣苔为中国特有的濒危植物,一度被认为灭绝后又被重新发现,引起了学界及公众的广泛关注.种子库是保护珍稀濒危物种的一种有效手段,对物种开展种子保存需要较为深入的了解其种子休眠及萌发特性;但对于弥勒苣苔,现有资料十分有限,本研究系统地探讨了弥勒苣苔种子的休眠状态及萌发习性.在对其种子进行X光透视后,发现其种子的胚发育完全,且基本充满整个种子,因此认为该种子不具有形态休眠.在多个变温及恒温条件下对新鲜种子进行萌发测试,发现在较高温度(高于20℃)下萌发率很高且萌发迅速,赤霉素及低温层积处理能够显著提高种子在较低温度下(低于20℃)的萌发率,因此可以判定其种子存在浅的生理休眠.种子在恒温下的萌发率比相应的变温下萌发率要高,可能是其对所处荫蔽环境的一种适应.硝酸钾对种子萌发有显著的促进作用,在野外环境下,硝酸钾有对生境空窗及较少竞争的指示作用.     

Tolerance and recovery responses of playa halophytes to light,salinity and temperature stresses during seed germination

Halogeton glomeratus (M. Bieb.) C.A. Mey., Lepidium latifolium Linn. and Peganum harmala Linn. are distributed in temperate salt playa habitats of Upper Hunza, Pakistan. Seeds were germinated under various salinity (0–500 mM NaCl), light (12 h-light:12 h-dark and 24 h-dark) and temperature (5/15, 10/20, 15/25, 20/30, and 25/35 °C, dark/light) regimes for 20 days to determine the optimal conditions for germination and recovery of seeds from these factors when exposed to less than optimal conditions. Seeds that failed to germinate in dark were transferred successively to 12 h-photoperiod, salinity to distilled water and from various temperature regimes to 20/30 °C, to determine the effect of these stresses and the ability of these seeds to recover respectively. Highest seed germination (H. glomeratus and L. latifolium: 100%; P. harmala: 80%) was obtained in non-saline control at 20/30 °C in 12 h-photoperiod, however, increase in salinity progressively inhibited seed germination. Seed germination of H. glomeratus and P. harmala was substantially inhibited and that of L. latifolium was prevented in dark. Salinity and dark treatments have a synergistic effect in inhibiting seed germination of all species. No seed of any species germinated at 5/15 °C; germination was substantially inhibited at 25/35 °C both for H. glomeratus and P. harmala while L. latifolium failed to germinate at 25/35 °C. Rate of germination also decreased with an increase in salinity at all temperature regimes but this effect was minimal at optimal temperature regime of 20/30 °C. After successive elimination of light, salinity and temperature stresses, final seed germination was identical to respective controls. The results indicate that seeds of these temperate halophytes could endure environmental stresses without losing viability and germinate readily when these stresses are removed. Under the extremely variable conditions of the playa habitat these species are highly opportunistic exploiting the windows of opportunity available during spring or early summer.     

Polyamine oxidase 3 is involved in salt tolerance at the germination stage in rice

Soil salinity inhibits seed germination and reduces seedling survival rate,resulting in significant yield reductions in crops.Here,we report the identification of a polyamine oxidase,OsPAO3,conferring salt tolerance at the germination stage in rice(Oryza sativa L.),through map-based cloning approach.OsPAO3 is up-regulated under salt stress at the germination stage and highly expressed in various organs.Overexpression of OsPAO3 increases activity of polyamine oxidases,enhancing the polyamine cont...     

种子休眠与萌发的分子生物学的研究进展

休眠与萌发是植物种子对环境变化的适应特征,受许多基因调控和环境因子的影响.利用数量遗传学方法(如QTL分析)和突变等手段已对休眠和萌发特性进行了深入的遗传学研究.近些年来,随着分子生物学的快速发展,种子休眠和萌发研究已经深入到分子水平.分子生物学技术的运用,特别是基因表达、基因组测序和以双向凝胶电泳及质谱分析为技术基础的蛋白质组学分析,已成为研究种子休眠和萌发的新工具和新方向.本文主要就利用分子生物学方法研究种子休眠与萌发的进展给予简要综述.     

Morphological and physiological responses of the halophyte, Odyssea paucinervis (Staph) (Poaceae), to salinity

In this study, salt tolerance was investigated in Odyssea paucinervis Staph, an ecologically important C₄ grass that is widely distributed in saline and arid areas of southern Africa. Plants were subjected to 0.2%, 10%, 20%, 40%, 60% and 80% sea water dilutions (or 0.076, 3.8, 7.6, 15.2, 22.8, and 30.4 parts per thousand) for 11 weeks. Increase in salinity from 0.2% to 20% sea water had no effect on total dry biomass accumulation, while further increase in salinity to 80% sea water significantly decreased biomass by over 50%. Morphological changes induced by salinity included reductions in the number of culms, leaves and internodes as well as decreases in internode length and leaf length:leaf width ratios. Carbon dioxide exchange, leaf conductance and transpiration decreased at salinities of 40% and higher, while quantum yield of photosystem II (PSII), electron transport rate (ETR) through PSII and intrinsic photosynthetic efficiency generally decreased at salinities of 60% and higher compared to 0.2% sea water. Concentrations of Na⁺ and Cl⁻ increased significantly with salinity increase in both roots and shoots. Na⁺/K⁺ ratios in the roots and shoots ranged from 0.66 to 3.28 and increased with increase in substrate salinity. The maximal rate of secretion at 80% sea water was 415 nmol cm⁻² d⁻¹ for Na⁺ and 763 nmol m⁻² d⁻¹ for Cl⁻ with high selectivity for these two ions. Predawn and midday ψ decreased with increase in salinity and were more negative than those of the treatment solutions. The concentration of proline increased with increase in salinity in both roots and shoots. The data clearly indicated that O. paucinervis is a highly salt-tolerant species that is morphologically and physiologically adapted to a saline environment.     

Enhanced germination under high-salt conditions of seeds of transgenicArabidopsis with a bacterial gene (codA) for choline oxidase

Arabidopsis thaliana was transformed previously with thecodA gene from the soil bacteriumArthrobacter globiformis. This gene encodes choline oxidase, the enzyme that converts choline to glycinebetaine. Transformation with thecodA gene significantly enhanced the tolerance of transgenic plants to low temperature and high-salt stress. We report here that seeds of transgenic plants that expressed thecodA gene were also more tolerant to salt stress during germination than seeds of non-transformed wild-type plants. Seedlings of transgenic plants grew more rapidly than those of wild-type plants under salt-stress conditions. Furthermore, exogenously applied glycinebetaine was effective in alleviating the harmful effects of salt stress during germination of seeds and growth of young seedlings, a result that suggests that it was glycinebetaine that had enhanced the tolerance of the transgenic plants. These observations indicate that synthesis of glycinebetaine in transgenic plantsin vivo, as a result of the expression of thecodA gene, might be veryuseful in improving the ability of crop plants to tolerate salt stress. The extended abstract of a paper presented at the 13th International Symposium in Conjugation with Award of the International Prize for Biology “Frontier of Plant Biology”     

Action spectra for light-induced germination in dormant lettuce seeds

Fluence response curves for red light-induced germination of thermodormant (TD) seeds of Lactuca sativa L. show two regions that differ in their light sensitivity. In the region of high sensitivity, the germination responses differ between seed batches and can be altered by dark storage or far red irradiation. Induction of germination in far red dormant (FRD) seeds requires far higher fluences. Action spectra for induction to 60% germination were determined for these various response types. Spectra for the regions of low sensitivity response are similar for TD and FRD seeds. In comparison, the action spectrum for the highly sensitive response in TD seeds is significantly shifted to longer wavelengths. Analogous differences exist in the action spectra for far red reversal of the red induced germination responses. Germination induction in the low sensitivity region shows repeated red-far red reversibility. Far red reversal of red induction in the high sensitivity region does not saturate even at the highest far red fluences available and requires increased red fluences for subsequent reinduction. A model quantitatively accounting for these observations is presented. It is pointed out that action spectra of processes involving photoreversible pigments with partly overlapping absorption spectra in general are not identical with the absorption spectra of the partners. They should depend upon the degree of phototransformation required to elicit a given physiological response. In the case of induction of lettuce seed germination the observed action spectra can be interpreted as reflecting different requirements for P _fr of the various response types. Our results do not necessitate the assumption of spectroscopically different forms of phytochrome in these seeds.Abbreviations TD thermodormant - FRD far red dormant - P phytochrome - P _r red absorbing form of P - P _fr far red absorbing form of P     

Effects of seed storage on germination of two succulent desert halophytes with little dormancy and transient seed bank

Seeds of both Salsola imbricata and Haloxylon salicornicum have high germination level and germination speed, and form a transient seed bank in nature. The impacts of storage period and condition on germination level and speed were assessed in the two species. Storage for three months significantly increased both germination level and speed of seeds stored under the different conditions, compared to that of fresh seeds. In both species, nine months storage did not affect germination percentage in cold storage seeds, but completely inhibited it in field seeds. Storage for longer time in room and warm temperatures resulted in significant reduction or complete inhibition in the germination of the two species, so this was more pronounced in H. salicornicum. Storage significantly increased germinate rate index of seeds stored in all conditions till 17 months in S. imbricata and till 12 months in H. salicornicum. In both species, fridge storage had little effects on final germination and germination speed of seeds incubated at the different temperatures, compared to fresh seeds. However, room temperature and warm storages significantly reduced final germination and germination speed at the different temperatures, so the reduction was more pronounced at 35 °C, especially in H. salicornicum.     

以色列二穗短柄草(Brochypodium distachyon)种子的休眠与萌发特性

对以色列3个群体的二穗短柄草(Brochypodium distachyon)种子的休眠与萌发进行了研究.结果表明:以色列二穗短柄草种子都具有休眠的习性,且不属于外源休眠(种壳休眠)类型;储藏温度和时间等因素对不同群体的二穗短柄草种子的休眠和萌发有着极显著的影响,高温(40℃)储藏可以加快种子后熟,打破种子的休眠;3个群体的二穗短柄草种子休眠深度不同,依次是:Sede Boqer>Yatk>Mt.carmel;且休眠深度与群体生长地年平均降雨量呈负相关关系.二穗短柄草的这些休眠和萌发特性可能是其长期与环境相互作用,在自然选择压力下进化形成的存活机制.     

Comparative salt tolerance in triticale,wheat and rye during germination

Summary Differences in salt tolerance of wheat and barley have been observed but knowledge of such differences in triticale (X Triticosecale Wittmack) cultivars is of potential importance. Effects of six salinity levels (0 to 2% NaCl) on germination of triticales Beagle and 6TA 131 in comparison to wheat and rye were determined at 15–20°C. Beagle triticale and rye showed similar trends in germination reduction as salinity increased from 0 to 1.5% NaCl and exhibited fairly high salt tolerance. However, 6TA 131 triticale and wheat showed a significant drop in germination even at 0.5% NaCl concentration and were more susceptible to salt injury.     

Dormancy in somatic embryos and seeds ofVitis: changes in endogenous abscisic acid during embryogeny and germination

Abscisic acid (ABA) in extracts of somatic embryos and seeds of Gloryvine (Vitis vinifera L.xV. rupestris Scheele) was measured by gas chromatography-mass spectrometry-selected ion monitoring using deuterated ABA, (±)-[C-3Me-²H₃]ABA, ([²H₃]ABA) as internal standard. The ABA content increased rapidly during embryogeny (0.035 ng/embryo at the globular stage to 0.22 ng/embryo at the mature stage). The level of ABA in the tissues of somatic embryos, expressed in ng/mg dry weight, decreased from the globular stage (0.76 ng/mg) to the mature stage (0.25 ng/mg). Chilling (4° C) induced normal germination of seeds and mature somatic embryos and precocious germination of globular, heart-shaped and torpedoshaped somatic embryos. In all cases chilling led to a marked reduction in endogenous ABA. Exogenous (±)-ABA inhibited the germination of chilled somatic embryos.Abbreviations ABA abscisic acid - [²H₃]ABA (±)-[C-3Me-²H₃]-abscisic acid - BHT 2,6-di-t-butyl-4-methylphenol - GC-MS gas chromatography-mass spectrometry - Me-ABA and Me-[²H₃]ABA methyl esters of ABA and [²H₃]ABA, respectively - SIM selected ion monitoring     

Endo-β-mannanase activity during dormancy alleviation and germination of white spruce (Picea glauca) seeds

It is not known how embryos of seeds of the Pinaceae protrude from their enclosing tissues to complete germination. Prior to protrusion of the radicle there is an increase in endo-β-1,4-mannanase (EC 3.2.1.78) activity associated with weakening of the micropylar megagametophyte/nucellus from seeds of white spruce ( Picea glauca [Moench.] Voss). Mannanase activity is present as three isoforms (pI values 5.0, 4.8, 4.7) in both the embryo and surrounding structures (megagametophyte and nucellus) prior to and during imbibition. Activity of all the isoforms increases in the chalazal and micropylar megagametophyte during germination. Activity then declines after the testa splits, typically 1 day prior to radicle protrusion, due partially to its leaching from the seed into the surrounding water. Activity increases in the cotyledons and axis as the embryo commences elongation. Seeds from dormant seedlots exhibit a lower germination percentage, relative to seeds from nondormant seedlots, and the force necessary for the embryo to puncture the surrounding structures tends to be greater. Although similar mannanase activities are present in unimbibed seeds of dormant and nondormant seedlots, during germination, enzyme activity in seeds of dormant seedlots is lower. Moist chilling alleviates dormancy in the seeds of the Pinaceae and, during 3 weeks of this treatment, mannanase activity slowly increases. After 3 weeks of moist chilling and regardless of whether the seedlot was dormant or not prior to moist chilling, the force necessary to puncture the micropylar megagametophyte and nucellus is lower, and the speed of germination greater. Seeds from previously dormant seedlots also complete germination to a greater percentage, relative to unchilled seeds from dormant seedlots. Upon transfer to 25°C, mannanase activity in moist-chilled seeds decreases during germination of all seedlots regardless of their previous dormancy status.     

Dormancy and germination of olive embryos as affected by temperature

Olive seeds do not germinate promptly when placed under favourable conditions, which is a problem in raising young plants for breeding or experimental purposes. In a series of experiments an investigation of the role of temperature in the germination of olive embryos was conducted. Naked, unchilled olive embryos ( Olea europaea L. cv. Chalkidikis), cultured in vitro at 20°C, had a germination capacity of 73%, whereas that of embryos which had previously been chilled at 10°C for 2 or more weeks reached 96%. Intact seeds did not germinate at 20°C unless they had previously been subjected to 10°C for 3 or 4 weeks. Embryos chilled while in the intact seed and excised just before transfer to 20°C, reacted in a similar way to naked embryos, but reached their maximum germination capacity after 4 weeks at 10°C. Under constant temperature conditions the highest germination percentage of embryos was observed at 10 and 15°C and the highest germination rate at 15°C, while a moderate capacity and rate of germination occurred at 20°C, and a very low percentage and rate at 25 and 30°C. Prechilling at 10°C did not affect germination at 15°C, but improved the percentage and the rate of germination at 20, 25 and 30°C. The germination percentages of embryos chilled for 1 or 2 weeks at 10°C and then transferred to 25°C were lower than those of similarly chilled embryos transferred to 20°C. The chilling effect could not be reversed at 25°C when the embryos had been chilled for 3 or more weeks. The results show that olive seeds exhibit a state of dormancy that is caused by factors residing partly in the endosperm and partly within the embryo.     

Dormancy inDioscorea: Differences of temperature responses in seed germination among six Japanese species

Effects on seed germination of temperatures ranging from −2 C to +29 C were tested inDioscorea nipponica, D. tokoro, D. japonica, D. tenuipes, D. septemloba andD. quinqueloba which orginate in the temperate zone; they are distributed from northern cold areas to southern warm areas approximately in this order in Japan. After water imbibition of these seeds, chilling induced full germination, and high temperatures over 23 C induced a secondary dormancy, but sensitivities to the chilling and to the high temperatures differed with species. Cold-climate species germinated rapidly at higher temperatures after a short-term chilling or even without chilling, whereas warm-climate species required chilling of a rather long period for germination; thus, among 6 species tested, favourable temperatures for germination and climatic temperatures of distribution area were conversely correlated. Seeds ofD. tokoro andD. japonica collected from several populations grown in different climates were also tested for germination at 11 to 29 C; seeds from warm climates germinated rather slowly compared to seeds from cold climates. These inte- and intra-specific adaptation manners in the temperature members of the genusDioscorea are entirely different from those of many other plant genera reported by some workers.     

Growth physics and water relations of red-light-induced germination in lettuce seeds

Red light (R) and gibberellins (GA) each induce a water potential decrease in the axes of lettuce (Lactuca sativa L.) embryos resulting in germination of intact seeds (achenes) or an increase in growth of the axes of isolated embryos. The fruit coat and endosperm are a substantial barrier to the penetration of exogeneous GA. Isolated embryos take up 35 times as much [³H]GA₁ as the embryos of intact seeds and respond to less than 1·10^-10 M GA₃ or GA₄₊₇. We calculated that only 1·10^-8 M of either GA₃ or GA₄₊₇ would result in 50% germination if the GA were able freely to penetrate the fruit coat. Exogenous GA₃ or GA₄₊₇, at concentrations insufficient to cause germination, result in an apparent synergistic promotion of germination when suboptimal R is applied. Yet suboptimal concentrations of exogenous GA₃ or GA₄₊₇ and suboptimal R result in only additive increases in the growth response in axes of isolated embryos. Dose-response curves demonstrate quantitative increases in the growth response of the isolated axes after R or GA treatments insufficient to induce germination in intact seeds, indicating that a threshold potential must be achieved by the embryonic axes before germination can occur.Abbreviations FR far=red light - GA gibberellin - PEG poly-ethylene glycol 4000 - P_fr far-red-absorbing phytochrome - R red light III.=Carpita et al. 1979b; IV.=Carpita et al. 1979c     

箭叶淫羊藿种子的休眠类型与萌发研究

箭叶淫羊藿具有重要的药用价值和较大的市场需求,但其种子休眠和萌发特性目前仍不够清楚,严重影响了其产业化育苗与栽培。为探明其种子的休眠类型及释放休眠和促进休眠的最适方法,该文以箭叶淫羊藿的成熟种子为材料,研究种子对水分的吸收、种子的脱水耐性,以及温度、层积和植物激素对种子休眠与萌发的影响。结果表明：(1)箭叶淫羊藿种子不具有物理休眠,而具耐脱水性。(2)在4～25℃和黑暗条件下,种子萌发率为零,具有休眠特性。(3)胚与种子的比值非常小,4℃和10℃及变温层积能显著地促进胚的生长发育以及增加种子的萌发速率和萌发率。(4)赤霉素和氟啶酮显著增加种子的萌发速率和萌发率。综上认为,箭叶淫羊藿种子的休眠类型为形态生理休眠(MPD),释放休眠和促进萌发的最适方法是先将种子在10℃中层积30 d,再置于4℃环境中让其萌发。该研究结果为箭叶淫羊藿的产业化育苗提供了参考。