A high temperature requirement for after ripening of imbibed dormant Poa annua L. seeds

Freshly harvested seeds of Poa annua L. collected in south Louisiana were stored in moist soil at seven temperatures between 5°C and 35°C. At monthly intervals, seed lots were removed and germinated at each of the seven temperatures. Seed were dormant for at least 1 month at all test temperatures. Seeds stored for 2 months at 30 and 35°C showed conditional dormancy; there was 100% germination at 10 or 15°C, and poorer germination at 5 or 20°C. Seeds started to lose viability after 2 months at 35°C and were dead after 7 months. In seeds stored at 10–30°C, there were increased percentages and a wider range of germination temperatures as storage time or storage temperatures increased. Seeds stored at 10°C remained dormant for 9 months, but by 12 months of storage the seeds germinated only at 5 or 10°C. Nearly all seeds stored at the same temperatures in air dry soil remained dormant for 6 months, regardless of storage temperature. These results differ from other reports of low temperatures breaking seed dormancy in Poa annua L. and suggest an adaptation to subtropical climates.     

Dormancy studies in three populations of Poa annua L. seeds

Seeds of Poa annua from original collections in Louisiana, Maryland and Wisconsin were grown together in Louisiana over a 3-year period. The freshly harvested seeds and samples stored in moist soil at 30°C were tested for germination at a range of temperatures to compare dormancy and germination characteristics. Seeds of the Louisiana population were dormant over the germination temperature range of 5–25°C, and imbibed storage for 2 weeks did not break dormancy. Freshly harvested seeds of the Maryland population germinated well (78%) at 10°C. With 1 week of imbibed storage at 30°C, germination was good over the range from 5 to 15°C and near 50% at 20°C. Storage for 2 weeks had little further effect. Freshly harvested seeds of two Wisconsin populations germinated above 50% throughout the range of temperatures, and imbibed storage for 2 weeks at 30°C had no effect on germination. The variations in the dormancy of freshly harvested seeds and the varying responses of dormancy breaking from storing imbibed seeds at 30°C suggests that these populations have adapted to avoid high summer temperatures in Louisiana and Maryland but to grow as a summer annual in Wisconsin.     

Effect of temperature during burial on dormant and non-dormant seeds of Lamium amplexicaule L. and ecological implications

Spring-produced seeds of Lamium amplexicaule L. were dormant at maturity in May and after-ripened when buried and stored over a range of temperatures, becoming conditionally dormant at low (5, 15/6 and 20/10°C) and non-dormant at high (25/15, 30/15 and 35/20°C) temperatures. Conditionally dormant seeds germinated to high percentages at 5 and 15/6°C, and non-dormant seeds germinated to high percentages at 5, 15/6, 20/10, 25/15 and 30/15°C. Seeds that became conditionally dormant at 5°C afterripened completely (i.e. became non-dormant) after transfer to 30/15°C. Buried seeds that became non-dormant in a non-temperature-controlled glasshouse during summer were still non-dormant after 12 weeks of storage at 30/15°C, while those stored at 5°C for 12 weeks had entered conditional dormancy. Thus, low temperatures cause reversal of the afterripening that takes place at high temperatures, but not that which takes place both at low and at high temperatures. Low winter temperatures cause dormant autumn-produced seeds and non-dormant seeds in the soil seed pool to become conditionally dormant. The ecological consequences of these responses to temperature are discussed in relation to the timing of seed germination in nature.     

Role of temperature in regulating timing of germination in soil seed reserves of Lamium purpureum L.

Fresh seeds of Lamium purpureum L. were dormant at maturity, and when buried and exposed to natural seasonal temperature changes they exhibited an annual dormancy/non-dormancy cycle. During burial in summer, fresh seeds and those that had been buried for 1 year afterripened and thus were non-dormant by September and October; light was required for germination. During autumn and winter seeds re-entered dormancy, and during the following summer they became non-dormant again. Dormant seeds afterripened when buried and stored over a range of temperatures, becoming conditionally dormant at low (5, 15/6°C) and non-dormant at high (20/10, 25/15, 30/15 and 35/20°C) temperatures. Conditionally dormant seeds germinated to high percentages at 5, 15/6 and 20/10°C, while non-dormant seeds germinated to high percentages additionally at 25/15, 30/15 and 35/20°C. Low temperatures caused non-dormant seeds to re-enter dormancy, while high temperatures caused a sharp decline in germination only at 30/15 and 5°C. The temperature responses of L. purpureum seeds are compared to those of L. amplexicaule L.     

Dormancy and viability of Phalaris minor seed in a rice–wheat cropping system

Seed placement, soil temperature and soil moisture content influenced the process of after-ripening in Phalaris minor seeds. Seeds of P. minor collected from the soil just after wheat harvesting exhibited higher germination than seeds from P. minor threshed directly. There was a pronounced impact of periodic inhabitation of seed into the soil on germination after its dispersal. Germination was strongly inhibited when the seed was kept in soil at more than field capacity (FC) or in water. Maximum germination of seed incubated in soil at FC occurred at 30°C while a temperature of 40°C favoured after-ripening of seed when mixed with dry soil or kept dry without any medium. Release from conditional dormancy was quicker in the seed retrieved from the soil kept at 20°C than at 10°C. Seed release from conditional dormancy and germination increased with a rise in temperature from 30 to 40°C when the seed was retrieved from incubation in soil at FC for 70 days. The seed kept immersed in water was least responsive to a rise in temperature. Seed recovered from dry soil, or kept without any medium, responded quickly at both temperatures. Light enhanced the germination of Phalaris minor seed. The seedbank subjected to rice (Oryza sativa) field management conditions lost vigour in comparison with the seed stored in laboratory. There was significant variability in seed viability when exposed to differential water management conditions in rice.     

Field emergence and temperature requirements for germination in Solanum sarrachoides Sendt.

Emergence of Solanum sarrachoides began in late April, reached a peak in May or June and ceased in September. This pattern closely resembled that for S. nigrum L., whereas almost all seedlings of S. dulcamara L. emerged in April. Fresh seeds of S. sarrachoides were dormant but developed a capacity for germination at 25 and 30°C and at alternating (16 h low/8 h high) temperatures of 4/25, 10/25, 10/30 and 20/30°C when stored dry. kept moist at 4°C or buried in the field. Buried seeds also became capable of germinating at 10. 15 and 20°C and the temperature range for germination was widest during April-June. Induced dormancy developed during August and the range narrowed. The consistent seasonal emergence pattern appears to be associated with cyclic changes in the dormancy status of buried seeds.     

Germination responses of buried seeds of Capsella bursa-pastoris exposed to seasonal temperature changes

Buried seeds of Capsella bursa-pastoris exhibit an annual conditional dormancy/non-dormancy cycle. Seeds after-ripen during summer and remain non-dormant during autumn and winter. Seeds enter conditional dormancy in early spring, first showing marked decreases in ability to germinate at high (35/20°C) and then at lower (30/15, 25/15°C) temperatures. Seeds do not lose the ability to germinate to high percentages at March (15/6°C) and April (20/10°C) temperatures in March and April. Thus, C. bursa-pastoris is a facultative winter annual, germinating in both autumn and spring if seeds are exposed to light. However, because some seeds retain the ability to germinate at 30/15 and 25/15°C, they could do so throughout the growing season in regions with cool, moist summers. Conditional dormancy developed in all seeds given 12 weeks at 5°C and subsequently kept for 4 weeks each at March (15/6°C), April (20/10°C) and May (25/15°C) temperatures. Thus, seeds of C. bursa-pastoris enter conditional dormancy as temperatures increase in spring.     

Germination ecology of seeds of the annual weeds Capsella bursa-pastoris and Descurainia sophia originating from high northern latitudes

Low temperatures may inhibit dormancy break in seeds of winter annuals, therefore it was hypothesized that seeds of Capsella bursa‐pastoris and Descurainia sophia that mature at high latitudes in late summer–early autumn would not germinate until they had been exposed to high summer temperatures. Consequently, germination would be delayed until the second autumn. Most freshly matured seeds of both species collected in August and September in southern Sweden were dormant. After 3 weeks of burial at simulated August (20/10°C) and September (15/6°C) temperatures, 28 and 27%, respectively, of the C. bursa‐pastoris and 56 and 59%, respectively, of the D. sophia seeds germinated in light at 15/6°C. In contrast, in germination phenology studies conducted in Sweden, only a few seeds of either species germinated during the first autumn following dispersal. However, there was a peak of germination of both species the following spring, demonstrating that dormancy was lost during exposure to the low habitat temperatures between late summer and early autumn and spring. Nearly 100% of the seeds of both species subjected to simulated annual seasonal temperature changes were viable after 30.5 months of burial. In the burial study, exhumed seeds of C. bursa‐pastoris were capable of germinating to 98–100% in light at the simulated spring–autumn temperature regime (15/6°C) in both spring and autumn, while those of D. sophia did so only in autumn. In early spring, however, seeds of D. sophia germinated to 17–50% at 15/6°C. Thus, most seeds of these two annual weeds that mature in late summer do not germinate in the first autumn, but they may do so the following spring or in some subsequent autumn or spring.     

Longevity,dormancy and germination of Cyanus segetum

Cyanus segetum is an iconic, colourful weed in arable fields that provides ecological and societal services. To understand better both the infestation dynamics of C. segetum as an abundant, harmful weed and maintain sustainable populations where it provides beneficial services, we compared information on seed dormancy, seed longevity and germination conditions in two populations. Persistence of seeds buried in the soil was low, with <10% viable after 3 years. Periodic dormancy cycling was observed over the 4 years in the soil, with a maximum of dormant seeds in the spring and a minimum in the autumn; however, 20% of the seeds were non‐dormant all the time. Seeds of C. segetum were positive photosensitive, but light requirement varied among populations. Base water potential for germination was ?1 MPa. Base temperature ranged from 1 to 2°C. Optimum temperature for germination was about 10 to 15°C, but the mean thermal time varied greatly between populations, from 80 to 134 day °C. Photoperiod and temperature combinations had no effect on germination percentage, but both reduced the germination rate. Burial deeper than 2 cm greatly reduced germination and seedling emergence strongly decreased at depths >0.5 cm. No seeds buried deeper than 8 cm emerged. Low seed longevity and a wide range of germination conditions could partly explain the rapid disappearance of C. segetum populations after herbicide application began in western Europe. However, yearly sowing in restoration areas does not seem to be essential.     

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.     

Dormancy,germination and emergence of Urochloa panicoides regulated by temperature

Urochloa panicoides is an annual weed of summer crops. In Argentina, in subhumid areas with monsoon rainfall, it germinates and establishes in a single flush. To (i) identify the environmental factors that modify its seed dormancy level and germination and (ii) quantify the parameters describing the thermal behaviour of the germination and emergence dynamics of this weed under non‐limiting water conditions, we established a set of germination experiments performed (i) under controlled conditions using seeds after ripened for 3 or 6 months in different thermal and hydric conditions and (ii) under field conditions, where the soil temperature was modified by applying different shading levels. Seed dormancy level remained high with 3 months after ripening in all treatments. After 6 months, seeds stored at 4°C in dry conditions did not germinate at any temperature, while seeds stored at 25°C in dry conditions and in situ germinated c. 20% and 60% respectively. Germination percentage was higher in seeds harvested before their natural dispersal. The base, optimum and maximum temperatures for seed germination were 6, 35 and 45°C respectively. Shading reduced the number of emerged seedlings, possibly by reducing the soil thermal amplitude. The results explained the dormancy‐breaking mechanism of U. panicoides that allows a high germination rate in the field when rainfall occurs.     

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.     

Germination ecology, emergence and host detection in Cuscuta campestris

Trials were carried out to study the germination and dormancy of Cuscuta campestris Y. (dodder) seeds and factors influencing the success of early parasitisation of sugarbeet. Primary dormancy can be removed by seed scarification. Germination was negligible at 10°C and optimal at 30°C, while it was not influenced by light. Seed burial induced a cycle of induction and breaking of secondary dormancy. Seedling emergence was inversely proportional to the depth of seed burial and only seed buried within 5 cm of the soil surface emerged. Storage of C. campestris seeds in a laboratory for 12 years resulted in the loss of primary dormancy, enabling the germination of all viable seeds. Host infection (i.e. protrusion of parasite haustoria from host tissue) was heavily influenced by host growth stage. Tropism towards a host was due to the perception of light transmitted by green parts of sugarbeet plants. Insertion of a transparent glass sheet between host leaves and parasite seedlings did not modify this response. This phototropism permitted Cuscuta to identify host plants with high chlorophyll content as a function of the lower red/far red ratio of transmitted light.     

Temperature requirements for after-ripening in buried seeds of four summer annual weeds

Freshly matured, seeds of the four summer annuals Ambrosia artemisiifolia, Polygonum pensylvanicum, Amaranthus hybridus and Chenopodium album were buried in soil at (12/12 h) daily thermoperiods of 15/6, 20/10, 25/15, 30/15 and 35/20°C and at a constant temperature of 5°C. After 0, 1, 3 and 5 months, seeds of each species at each temperature were exhumed and tested at a 14-h daily photoperiod at all six temperatures. Fresh seeds of A. artemisiifolia and P. pensylvanicum did not germinate at any temperature, those of A, hybridus germinated to 4 and 64% at 30/15 and 35/20°C, respectively, and those of C. album to 11–20% at 25/15, 30/15 and 35/20°C. Seeds of A. artemisiifolia and P. pensylvanicum, which germinate only in spring, required exposure to low (5, 15/6°C) temperature to after-ripen completely (i.e., to gain the ability to germinate over a wide range of temperatures), and little or no after-ripening occurred at high (25/15, 30/15 and 35/20°C) temperatures. Seeds of A. hybridus and C. album, which germinate in spring and summer, required exposure to low temperature to after-ripen completely, but at high temperatures they rapidly gained the ability to germinate at high temperatures. Regardless of the burial temperatures and species, when after-ripening occurred, seeds firs germinated at high and then at low temperatures. The minimum germination temperature for a species decreased with after-ripening temperature and with an increase in the length of the burial period.     

Seed dormancy and periodicity of seedling emergence in Veronica hederifolia L.

Emergence of Veronica hederifolia seedlings began in mid-October and continued into spring; few appeared from June to September. Ripe seeds shed in June were dormant but wben buried in soil outdoors developed a capacity for germination initially at low temperatures (constant4 C; daily alternations of 4-10° and 4-1 5 C) and later at somewhat higher temperatures, with peak germination in September-November. During winter, spring and early summer thc germination capacity declined, to increase again in late summer and early autumn. Cyclic physiological changes thus occur in seeds of V,hederifolia present in the soil, with which lhe consistent seasonal periodicity of seedling emergence is associated. In dry storage ihe capacity for germination progressively increased, but alter 12 months there was a sharp decline in germination at 4° C. Few seeds germinated at 20° C, but moistening with GA 4/7; brought about complete germination at this temperature.     

Seasonal changes in the germination responses of buried Lamium amplexicaule seeds

Spring-produced seeds of Lamium amplexicaule L. were buried in pots of soil in an unheated glasshouse in June 1978, and at 1–2-month intervals, for 27 months, they were exhumed and tested for germination in light and darkness at temperatures simulating those in the habitat from early spring to late autumn. Freshly-matured seeds were dormant, but by autumn 85% or more germinated in light at 15/6, 20/10, 25/15 and 30/15°C but only 7% or less in darkness. During late autumn and winter germination in light decreased at 25/15 and 30/15 °C but not at 15/6 and 20/10 °C, and germination in darkness increased at 15/6 and 20/10 °C. During late winter and early spring germination in light at 15/6 and 20/10 °C decreased, and seeds lost the ability to germinate in darkness. By the second autumn of burial, seeds germinated to near 100% in light at 15/6 to 30/15 °C and to 10–25% in darkness at 15/6 and 20/10 °C. The cycle of germination responses was repeated during the second winter and spring and the third summer of burial. Autumn-produced seeds were dormant when buried in November 1979, but by spring they germinated to 81 and 36% at 15/6 and 20/10 °C, respectively, in light. These seeds afterripened further during summer. The consequence of seasonal changes in germination responses is that (1) seeds can germinate in the habitat in late summer, autumn and spring but not in early- to mid-summer or in late autumn and winter and (2) during both germination seasons, seeds produced during the previous spring(s) and/or autumn(s) can germinate.     

Modelling Chloris virgata germination and emergence under different temperature and light quality conditions

Chloris virgata is a problematic weed around the world. Prediction of weed germination rates could be a useful strategy to optimise timing of weed control actions. We studied the germination and emergence of C. virgata collected seeds under different after-ripening treatments and different exhumation dates after seed dispersal, to estimate seed dormancy level and predict weed emergence dynamics under field conditions. Three experiments were conducted under controlled conditions to determine base, optimum and maximum germination temperatures (T_b, T_o and T_m respectively) and comprised: (a) exposure of seeds to gradually increasing and decreasing temperatures between 5 and 35°C; (b) exposure of seeds to different constant temperatures; and (c) exposure of seeds to different light quality conditions (red – far red ratio) and temperature regimes (constant and alternating temperatures). To explore genuine environmental conditions, a field experiment was performed to determine weed emergence under different shading levels. Finally, with the data obtained, a thermal time model for dormancy release was used to predict C. virgata seedling emergence in the Argentine Pampas region. Seeds after-ripened in cold and wet conditions and constant 25°C showed the highest germination percentages. The values of T_b (7°C), T_o (28°C) and T_m (40°C) remained constant at all exhumation dates. Neither light quality nor thermal regime modified the final germination percentages. However, shading delayed seedling emergence under field conditions, even when it was adjusted by thermal time. These results may allow predicting C. virgata emergence in temperate regions and help to improve weed control in integrated weed management strategies.     

Asian spiderflower (Cleome viscosa) germination ecology in southern Iran

Cleome viscosa is one of the most important weeds of warm‐season crops in southern Iran. Laboratory experiments were conducted to assess the impact of environmental factors on seed germination of C. viscosa . Freshly harvested seeds exhibited dormancy that was relieved (>90%) after immersion for 20 min in concentrated sulfuric acid. Regardless of the temperature regime, the final percentage of germination in light/dark (69.3%) was significantly higher than in complete darkness (58.3%). The optimum temperature for germination was 35/25°C in both light and dark. No germination was observed at constant temperatures of either 15 or 45°C. The thermal thresholds for seed germination, the base (T _b) and the mean ceiling germination temperatures (T _c(50)) were estimated to be 18.8 and 39.9°C, respectively. A base water potential ( Ψ _b(50) ) of ?0.96 MPa was identified for C. viscosa seeds. The response threshold of C. viscosa to reduce 50% of maximum germination for salinity was estimated to be 255 mM. Seeds that were placed on the soil surface had the highest percentage of seedling emergence (77.3%), and no seedlings emerged from seeds placed at a depth of 6 cm. The findings of this study could help to improve the integrated weed management strategies for this species.     

Study of seasonal variation in dormancy of Spergula arvensis L. seeds in a condensed annual temperature cycle

Changes in dormancy of Spergula arvensis seeds were studied during pre-incubation at constant temperatures and under a temperature regime that condensed the annual temperature cycle into 73 days. Each day in the regime represented the mean day and night temperatures and day lengths of 5 successive days of an average year in The Netherlands. Incubation occurred in water or loamy sand, in darkness. Germination of the seeds was tested in water or KNO₃ over a range of temperature. Seeds were irradiated with saturating doses of red light. In half of the treatments, pre-incubated seeds were dehydrated at the transfer to the conditions of the germination test. Breaking of dormancy occurred under conditions of 'spring'. It did not depend on exposure to low‘winter’temperatures, but was induced by rising 'spring’temperatures. Seeds developed secondary dormancy in late‘autumn'. The expression of the changes in dormancy that were induced during pre-incubation depended on the conditions of the germination test. Light, nitrate and dehydration stimulated germination. The experiments predicted that field emergence from nitrate-poor soils that have not been dehydrated will be restricted to a short period in autumn, whereas disturbance of nitrate-rich soils followed by a dry spell will stimulate germination of S. arvensis seeds from early spring to late autumn. The data presented good explanations for the cosmopolitan character and the serious weediness of this species. Its classification as a summer or winter annual is discussed.     

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.