Germination strategies of annual and short-lived perennial species in the Arabian Desert

Germination timing is highly regulated in short-lived plant species since it strongly influences recruitment success of vegetation. In deserts, the spatiotemporal distribution of plant-available water is highly episodic and unpredictable, making winter months more favorable for seed germination when other abiotic conditions co-occur. We hypothesized that changes in photoperiod and thermoperiod would impact germination more in seeds that had undergone in situ storage. We assessed 21 annual and short-lived perennial species in the Arabian Desert to find (1) if seeds were dormant at maturity, (2) if in situ seed storage increased germination percentage compared with no storage, (3) if photoperiod and thermoperiod germination requirements were influenced by in situ storage, and (4) if a phylogenetic association in seed germination could be observed. Seeds of each species collected in early 2017 were divided into two batches. One was tested for germination within one week (fresh seeds). The other was stored in situ at the maternal location (stored seeds) until October 2017 and tested for seed germination in the first week of November. Seed germination was conducted in incubators at two thermoperiods (15°C/20°C and 20°C/30°C; 12 h/12 h), and two photoperiods (12 and 0 h light per day). Results indicated that seed germination percentages of 13 species were significantly enhanced by in situ storage. A thermoperiod response was exhibited by stored, but not fresh seeds. Light exposure increased germination of fresh seeds but had only a minimal effect on stored seeds. Germination traits exhibited no phylogenetic correlation. This result indicated that selection pressure for germination strategy was stronger than that for taxonomic traits of these desert species.     

温周期、贮藏时间和盐分对短毛柽柳种子萌发的影响

短毛柽柳(Tamarix karelinii)是分布于盐渍化沙地和重盐碱地的灌木。为了阐明短毛柽柳种子的萌发特性,设置不同的温周期、储藏时间和盐分梯度,采用室内控制实验研究这些因子对种子萌发的影响。结果表明:1短毛柽柳种子在4个温周期中均具有高的萌发率,并且温周期温度越高,种子的萌发速率越快。2室温贮藏1个月显著提高种子的萌发速率,在贮藏6个月内,种子的最终萌发率都能达到100%。3≤0.8 mol·L~(-1)的Na Cl溶液对种子的最终萌发率无显著影响,而≥1.0 mol·L~(-1)的溶液则抑制种子的萌发,且随着浓度的增加,种子的最终萌发率呈下降趋势,直至为0。将1.0~4.0 mol·L~(-1)溶液中没有萌发的种子,转移至蒸馏水继续培养后,仍有71.67%~28.00%的种子恢复萌发率,表明种子具有较强的耐盐性。对短毛柽柳种子萌发特性的研究,为短毛柽柳的种苗培育提供基础资料。     

Seasonal changes in germination responses of buried seeds of the weedy summer annual grass Setaria glauca

Seeds of Setaria glauca (L.) Beauv. buried in soil and exposed to natural temperature cycles exhibited seasonal changes in temperature, but generally not light; dark requirements for germination. Seeds were dormant at maturity in late September and October (autumn), and during burial from October to January they entered conditional dormancy, germinating up to ≥60% in light and darkness at daily thermoperiods of 25/15,30/15 and 35/20^C by January. During burial from February to May or June, seeds became non-dormant and germinated up to 68–100% in light and darkness at 15/6,20/10,25/15,30/15 and 35/20^C in May or June. At maximum yearly temperatures in June or July–August, 65–89% of the seeds entered conditional dormancy (germinating at 30/15 and 35/20, but not at 15/6,20/10 and 25/15^C), and the others entered dormancy (not germinating at any thermoperiod). Thus, most buried seeds had an annual conditional dormancy/non-dormancy cycle, but some had an annual dormancy/non-dormancy cycle. Except for seeds buried in 1990 that lost the ability to germinate in darkness at all thermoperiods the first summer of burial, seeds incubated in light and in darkness exhibited the same patterns of seasonal changes in germination responses. Although conditionally dormant and non-dormant seeds germinated to high percentages in darkness in Petri dishes, seedlings were found only in bags of seeds exhumed in April and May 1983, indicating that some factor(s) associated with the burial environment other than darkness prevented germination of buried seeds.     

Maternal salinity improves yield,size and stress tolerance of Suaeda fruticosa seeds

Shrubby seablite or lani(Suaeda fruticosa Forssk)is a perennial euhalophyte with succulent leaves,which could be planted on arid-saline lands for restoration and cultivated as a non-conventional edible or cash crop.Knowledge about the impacts of maternal saline environment on seed attributes of this important euhalophyte is lacking.This study investigated the effects of maternal salinity on yield,size and stress tolerance of S.fruticosa seeds.Seedlings of S.fruticosa were grown in a green net house under increasing maternal salinity levels(0,300,600 and 900 mM NaCl)until seed production.Total yield,size,stress tolerance and germination of the descended seeds under different maternal saline conditions were examined.Plants grown under saline conditions(300,600 and 900 mM NaCl)produce a substantially higher quantity of seeds than plants grown under non-saline condition(0 mM NaCl).Low maternal salinity(300 mM NaCl)improves seed size.Seeds produced under all maternal salinity levels display a higher tolerance to low temperature(night/day thermoperiod of 10℃/20℃),whereas seeds produced under 300 mM NaCl maternal saline condition show a better tolerance to high temperature(night/day thermoperiod of 25℃/35℃)during germination.Seeds from all maternal saline conditions germinate better in the 12 h photoperiod(12 h light/12 h dark)than in the dark(24 h dark);however,seeds produced from low and moderate maternal saline conditions(300 and 600 mM NaCl)show a higher germination in the dark than those from control and high maternal saline conditions(0 and 900 mM NaCl).In general,maternal salinity is found to improve yield,size and stress tolerance of S.fruticosa seeds.     

The role of light and alternating temperatures on germination of Polygonum aviculare seeds exhumed on various dates

The annual dormancy cycle was investigated in buried seeds of Polygonum aviculare L. exposed to natural temperature changes in Lexington, Kentucky, U.S.A. Seeds were exhumed monthly from December 1984 to February 1987 and tested in light (14-h daily photoperiod) and continuous darkness at 12/12-h daily alternating temperature regimes of 15/6, 20/10, 25/15, 30/15 and 35/20°C. During autumn and winter, seeds became non-dormant, and in March 1985 they germinated to 95-100% at all thermoperiods in light and to 7-61% in darkness. Seeds remained non-dormant during spring but became more specific in their germination requirements in early summer. During July and August 1985, seeds germinated to 17-53% in light at 30/15 and 35/20°C but to 0-10% at all other test conditions. By September, about 65% of the seeds were dormant, but the others were able to germinate under the higher alternating temperatures in light. A similar seasonal cycle was recorded in the following year through to the spring of 1987. The results confirm the seasonal pattern of dormancy in this species (Courtney, 1968) but indicate that alternating temperatures combined with light are important in determining germination potential in P. aviculare.     

Inter-population variabilities in seed mass and germination of Panicum turgidum and Pennisetum divisum in the desert of Kuwait

Understanding variability in seed germination among populations is essential for planning an effective germplasm collection for restoration and conservation purposes.The knowledge of germination and dormancy patterns among populations of desert grasses is crucial for determining the potential of the species and populations to be used for restoration and conservation as well as forage production.Variability in seed germination of Panicum turgidum Forssk and Pennisetum divisum(Gmel.)Henr.in the desert of Kuwait was evaluated in different populations in May 2017.Experiment of seed germination(25 seeds and 4 replicates)was conducted for each population at night/day temperatures of 15℃/20℃and 20℃/30℃under the following light condition:continuous darkness or 12 h/12 h light/dark.Results showed that seed masses of both species strongly varied according to their seed provenances,and both species produced heavier seeds in population with a higher soil electrical conductivity.Seed germination percentage considerably varied between two species,and the variation in P.turgidum was greater(17%–49%)than that of P.divisum(72%–93%).Germination percentage in P.turgidum was greater at high temperature(20℃/30℃)than at low temperature(15℃/20℃).However,temperature regimes had no effect on germination percentage of P.divisum seeds.Mean germination time of both species exhibited significant inter-population variability.This result is especially relevant to assure the selection of the best population of each species and the regeneration success of the species.Besides this,inter-population variability also provides valuable information for enhancing our understanding of the mechanisms that regulate seed germination and how they might be related to seed provenance.     

Maternal salinity environment affects salt tolerance during germination in Anabasis setifera: A facultative desert halophyte

The effects of maternal salinity and light incubation on the salinity tolerance of the facultative halophyte Anabasis setifera during their germination stages were assessed. Seeds were collected from non-saline habitats in Egypt and saline habitats in the United Arab Emirates(UAE). The seeds of the two populations were germinated in 0, 100, 200, 400, 600 and 800 m M Na Cl, and incubated at 25°C/15°C in both 12-h light and 12-h darkness regimes and continuous darkness. Significantly more seeds germinated in the Egyptian population than in the UAE population. Salinity tolerance was significantly greater with the Egyptian population than with the UAE population, especially under the conditions of higher salinities. The difference in salinity tolerance between the seeds of two populations was attributed to their seed mass. In addition, germination was significantly faster for the Egyptian population than for the UAE population. Most of the saline treated seeds were able to recover their germination when transferred to distilled water, but this depended on their maternal salinity and light incubation. Recovery from higher salinities was significantly better for the seeds under darkness than for those under light in the UAE population, but the reverse was true for the seeds in the Egyptian population. The higher salinity tolerance for the A. setifera seeds from the non-saline Egyptian population and the lower salinity tolerance for the seeds from the saline UAE population cannot explain their natural distribution. Further studies about other possible roles, such as levels of different promoting and inhibiting phytohormones, are needed to understand the importance of salinity as an environmentally induced maternal effect.     

Influence of after-ripening environments on the germination characteristics and seed fate of Italian ryegrass (Lolium multiflorum)

Italian ryegrass ( Lolium multiflorum Lam.) is an exotic winter annual weed that recently has seriously infested wheat fields in central Japan. To understand the influence of tillage systems on the germination ecology and seedbank dynamics of naturalized Italian ryegrass, we compared the fate of the seeds that were after-ripened in the soil with that of the seeds on the soil surface, and then examined the germination characteristics of the seeds retrieved from both treatments by germination tests in both a light/dark (12 h photoperiod) condition and in constant darkness. The seedling emergence was greater initially for the seeds after-ripened on the soil surface than for the seeds in the soil. The component of emergent seeds after-ripened on the soil surface increased as time passed during the summer, whereas that of emergent seeds in the soil scarcely increased. This contrasting emergence was, in part, attributable to the ability of the seeds to germinate at higher temperatures in the light/dark condition than in constant darkness. These results suggest that untilled systems, which leave many seeds on the soil surface, might promote the rapid depletion of Italian ryegrass seeds through germination before wheat sowing in early winter. In contrast, seed burial by tillage might inhibit the germination of the seeds and form a large transient seed bank available to germinate during the wheat-growing season.     

Germination ecology of Emex spinosa and Emex australis,invasive weeds of winter crops

Emex spinosa and Emex australis are invasive dicotyledonous weeds. The effects of various environmental factors on the germination of these weeds were investigated under laboratory and glasshouse conditions. Germination response of both species was lower at warmer temperature, and maximum germination was recorded at 20/12°C (day/night). Light stimulated germination in both species, but considerable germination also occurred under darkness. More than 80% of E. spinosa seeds germinated at pH between 6 and 9, whereas E. australis seeds germination was considerably decreased at pH 9. Emex spinosa was fairly tolerant to salinity as compared with E. australis and germination (21%) of E. spinosa occurred even at 200 mm NaCl. Both species were sensitive to osmotic stress, but E. spinosa tolerated more osmotic stress than E. australis. Temperature above 20/12°C (day/night) and low osmotic potential increased time to start germination and mean germination time (MGT), as well as decreased germination index (GI) of both species. Darkness resulted in increased MGT and decreased GI in both species when compared with 10 h photoperiod. Salt stress strongly increased time to obtain 50% germination and reduced GI of both species. In both species, an increasing burial depth decreased emergence percentage and emergence index and increased time to start emergence, although some seed emerged even at 10 cm burial depth. It was concluded that both species can germinate over a wide range of environmental conditions. However, E. australis was more sensitive under adverse environmental conditions compared with E. spinosa. This information on germination ecology may aid in developing tools and strategies for management.     

不同环境因素对猪殃殃种子萌发及出苗的影响

为探讨猪殃殃种子萌发的适宜环境条件,通过室内萌发试验研究了温度、光照、pH、水势、盐分对猪殃殃种子萌发以及埋土深度对其出苗的影响。结果表明,猪殃殃种子萌发的适宜环境条件为温度10~20℃、光暗12h交替培养以及pH 4~10,萌发率可达到50%以上。猪殃殃种子对水势比较敏感,萌发率随着水势的降低而降低,当水势为0时,萌发率为50.50%,水势降低至-0.6 MPa时,仅有个别种子萌发,萌发率为3.50%,当水势为-0.7 MPa时,种子萌发完全受到抑制;猪殃殃种子对盐分胁迫有一定忍耐力,当NaCl浓度为0.16mol/L时,仍有少数种子萌发,萌发率为16.50%;适于猪殃殃种子出苗的埋土深度是1~2cm,出苗率达到49.50%。研究结果为猪殃殃的出苗调控和综合治理提供了依据。     

影响丰花草种子萌发和幼苗生长的主要环境因子

运用培养皿滤纸法和盆栽法,分别研究温度、光照和水分条件对热带常见杂草丰花草种子萌发和幼苗生长的影响。结果表明:丰花草种子萌发温度范围广泛(5~40℃),随温度升高,丰花草幼苗的鲜质量和生长长度均呈现先升高后降低的趋势;恒温下30℃为其种子萌发的最适温度,其发芽率、发芽指数、活力指数均相对较高,而平均发芽时间最短;25℃时生长的幼苗最健壮,其芽长、根长和芽鲜质量、根鲜质量均显著高于其他温度时的对应值。而变温条件更有利于丰花草种子萌发,12 h D/12 h L、5℃/15℃时发芽率达55.63%,15℃/25℃时发芽率90%,25℃/35℃时的种子萌发率、幼苗长度、幼苗鲜质量均最大。光照不是影响丰花草种子萌发的关键因子,但光照可能有利于促进丰花草幼苗生物量向根部分配,光暗交替(12 h D/12 h L)条件下丰花草幼苗生长情况相对最好(长度根冠比为1.05∶1;鲜质量根冠比为0.51∶1)。水分是决定丰花草种子能否萌发的关键因素,25℃、12 h D/12 h L条件下,在土壤最大持水量60%时丰花草种子萌发和幼苗生长状况均最佳。     

Assessment of daily heat pulse regimes on the germination of six Amaranthus species

In Mediterranean environments, thermal requirements for seed germination of Amaranthus species are met in spring. Nonetheless, seedlings of these species start emerging in the field from late winter, although the theoretical heat sum thresholds required for germination are not likely to be reached under field conditions. We hypothesised that soil thermal fluctuations can reduce the heat requirements of these species. Cardinal temperatures and thermal times of six Amaranthus species (A. albus, A. cruentus, A. deflexus, A. graecizans, A. retroflexus and A. viridis) were determined on non‐dormant seeds at constant temperatures ranging from 10 to 40°C. Subsequently, germination response to heat pulses was studied by imposing a thermal fluctuation regime of 32/8°C with two different thermoperiods of 3/21 and 6/18 h. In the two thermoperiods, exposure to 32°C was imposed for a different number of days: from 1 to 12 and from 1 to 6 heat pulses cycles in the 3/21 and 6/18 h thermoperiods respectively. Cumulative germination, germination rate and mean germination time were evaluated. Heat sum requirements and final germination percentage were affected by thermoperiod and number of thermal cycles. Amaranthus spp. germination was higher and faster when seeds were submitted to 6 h compared with 3 + 3 h of heat pulses. Our data showed that heat sum requirements for germination may change depending on the way in which varying temperatures are imposed on germinating seeds.     

Germination response of Phalaris paradoxa L. seed to different light qualities

The influence of different light regimes on the germination of Australian and English populations of Phalaris paradoxa L. (awned canary‐grass) seed was investigated to determine the impact of changing tillage practices on weed infestation. Seeds of all biotypes were highly viable, but differed in levels of innate dormancy (26–99%). In one experiment seed from a single Australian biotype, either enclosed in the spikelet glumes or having the spikelet glumes removed, were exposed to nine light treatments. Germination was stimulated by red and white light, but was inhibited by far‐red light. Time to 50% germination was less for seed enclosed in the spikelet glumes than for naked caryopses, although the final percentage of seed germinating when still enclosed in the spikelet glumes was significantly lower than for naked caryopses. In another experiment, six Australian and English biotypes with varying dormancy characteristics were exposed to eight light treatments. Red light did not stimulate germination in the deeply dormant biotype, however stimulated all other biotypes. Germination in darkness was below 20% in all biotypes except for one where germination was 51%. To overcome dormancy seeds were imbibed and placed in darkness at 16°C for either 7 or 14 days prior to exposure to red or white light for a single 15‐min period. Dormancy in all biotypes was overcome indicating that a period of burial may decrease the dormancy level and increase seed sensitivity to light. This increased light sensitivity suggests that exposure to light during tillage may stimulate germination in P. paradoxa seed.     

Biology of Commelina benghalensis L. in south-eastern Queensland. 2. Seed dormancy, germination and emergence

Germination of freshly harvested seeds of Commelina benghalensis L. varied from 0–3% for small aerial seeds, 20–35% for large aerial seeds and from 33% for small underground seeds to 90% for large underground seeds. Innate dormancy of all seed types was completely overcome by clipping the seed coat. Exposure to 90°C dry heat for 2 h was also effective in increasing germination of the three strongly dormant seed types. Optimum temperature for germination varied with the different seed types. Periods of likely major weed infestation from the four seed types were predicted using soil temperature data. Exposure to light increased germination but was not essential and underground seeds responded more to light than aerial seeds. Optimum depth of emergence for the four seed types was from 0 to 50 mm and there was a positive correlation between maximum depth of emergence and seed weight.     

Germination, emergence and seedling growth of Hypericum perforatum L.

Germination of new seeds (1–6 months old) of Hypericum perforation L. was restricted by high temperatures (16h/8h, 20/30°C), darkness and a chemical inhibitor in exudate from the capsule, whereas germination of old seeds (9 years) was only restricted by the inhibitor. The effect of the chemical inhibitor and high temperatures was overcome, respectively, by washing seeds in water and by reducing temperatures to constant 15°C. Calcium in solution from CaCO₃ and from three different soils did not prevent the germination of new or old seeds or of seeds collected from five different locations. There were differences in the germination characteristics and dormancy mechanisms of seeds collected from different localities, Restriction of the emergence of seedlings by a covering of > 2 mm of soil appeared to be due to lack of seedling vigour rather than to lack of light. Seedling growth was much slower than in other pasture species. Thus the requirements for germination of H. perforatum of low temperature and moisture to wash away the chemical inhibitor favour its establishment but the slow growth of its seedling restricts its emergence and renders it extremely susceptible to competition from other plants.     

Field assessment of thermal after-ripening time for dormancy release prediction in Lolium rigidum seeds

Dormancy release was studied in four populations of annual ryegrass (Lolium rigidum) seeds to determine whether loss of dormancy in the field can be predicted from temperature alone or whether seed water content (WC) must also be considered. Freshly matured seeds were after‐ripened at the northern and southern extremes of the Western Australian cereal cropping region and at constant 37°C. Seed WC was allowed to fluctuate with prevailing humidity, but full hydration was avoided by excluding rainfall. Dormancy was measured regularly during after‐ripening by germinating seeds with 12‐hourly light or in darkness. Germination was lower in darkness than in light/dark and dormancy release was slower when germination was tested in darkness. Seeds were consistently drier, and dormancy release was slower, during after‐ripening at 37°C than under field conditions. However, within each population, the rate of dormancy release in the field (north and south) in terms of thermal time was unaffected by after‐ripening site. While low seed WC slowed dormancy release in seeds held at 37°C, dormancy release in seeds after‐ripened under Western Australian field conditions was adequately described by thermal after‐ripening time, without the need to account for changes in WC elicited by fluctuating environmental humidity.     

Germination characteristics of Amaranthus quitensis as affected by seed production date and duration of burial

Thermal requirements for the germination of Amaranthus quitensis, a common annual weed in Argentina, were studied. In addition, temporal changes in dormancy from seeds produced at different times during the growing season were examined. For this second objective, thermal and light requirements for germination were tested in seeds buried at different depths, with or without crop residues. Base and optimum temperatures for germination rates were 12.8°C and 37°C respectively. At dispersal time, maximum percentage germination was 60–70% and this was generally recorded at 35°C/25°C in a 14-h photoperiod. Seed germination tended to increase in later seed collection dates. Seeds of A. quitensis showed seasonal changes in germinability in the soil. In winter, germination of retrieved seeds increased to over 90% until summer, after which there was a decrease until the following winter when germination was close to 40%. There were no differences in germinability between burial depths and crop residue levels. Germination requirements for alternating temperatures and light tended to disappear after burial. Initial viability was 99% and declined slightly during burial. Soil temperature seems to play a crucial role not only by regulating seasonal changes in dormancy, but also by defining the percentage and the germination rate in non-dormant seeds.     

Light, temperature and burial depth effects on Rumex obtusifolius seed germination and emergence

Trials were carried out to investigate the effects of light and temperature on germination of Rumex obtusifolius L. After several months of storage, seeds gradually lost dormancy and became photosensitive. Thermal optima for germination were between 20 °C and 25 °C in light or in darkness. At lower temperatures there was a greater demand for light, so that the greatest differences in germination percentage (between low and high temperatures) were found within the 10–15 °C temperature range. The calculated thermal minima ( x -intercept method) in light and darkness were 8.3 °C and 6.1 °C respectively. Daily temperature fluctuation increased germination even after seed irradiation with far-red light, suggesting a lower demand for the far-red-absorbing form of phytochrome. Seed burial inhibited germination in proportion to depth; however, germination inhibition was independent of seed phytochrome photo-equilibrium, which had been diversified by seed pretreatment with light. Seedlings did not emerge when seeds were buried >8 cm deep. Recovery of ungerminated seeds showed that excessive burial did not impede seedling emergence but rather prevented seed germination. However, this induction of dormancy was lost once germination processes were activated (24–48 h at 20 °C) that made germination irreversible. Temperature was also involved in inhibition, and low temperature (<15 °C) induced the least inhibition. This is discussed in terms of processes of respiration and fermentation in buried seeds.     

不同化学试剂及人工处理对稗草种子休眠的影响

分别应用不同化学试剂及人工剥去颖壳处理来提高稗草种子的萌发率,不同处理对种子发芽率的影响不同.GA3、浓H2 SO4、KNO3、剥去颖壳处理都有利于稗草种子的萌发,但剥去颖壳破除休眠的效果不理想.通过800～1200 mg/L GA3浸种24 h,浓硫酸浸种10～ 20 min,都能打破稗草种子休眠,使其发芽率超过70％;KNO3溶液可打破部分稗草种子休眠,2％ KNO3浸泡12 h,发芽率为34.67％;NaOH、HC1不宜用于解除稗草种子休眠.     

鹅观草种子萌发及出苗特性

在实验室和温室条件下研究不同温度、光照、水势、盐分对鹅观草种子萌发以及埋土深度对鹅观草出苗的影响。结果表明,鹅观草种子萌发的昼/夜温度范围为20℃/10℃~35℃/25℃,最适昼/夜温度为25℃/15℃;黑暗条件下,鹅观草种子萌发率较光照下略有提高,说明鹅观草萌发不需要光照;鹅观草种子对水势胁迫较敏感,水势从0下降至-0.8 MPa时,发芽率从93%直线下降至0;鹅观草种子对盐分胁迫具有一定耐受性,当NaCl浓度≤50 mmol/L时,种子均能保持80%以上的萌发率。50%萌发抑制率时的水势和NaCl浓度分别约为-0.4 MPa和100 mmol/L。埋土深度为0.5 cm时,鹅观草出苗率最高,埋土深度≥3 cm时出苗率显著下降。表明鹅观草种子萌发具有较强的环境适应性,通过耕作将其种子带入4 cm以下土层,可有效抑制鹅观草的危害。