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.     

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.     

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 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.     

Factors affecting seed germination and emergence of button grass (Dactyloctenium radulans) (R.Br.) P.Beauv.

Button grass (Dactyloctenium radulans) is a native, widely spread summer grass weed species in Australia. However, limited information is available on the seed germination biology of this species. Experiments were conducted to evaluate the effect of environmental factors on the germination and emergence of two populations of D. radulans. The seeds of these populations were collected separately from Dalby, Queensland and Coleambally, NSW. Seeds were germinated at a range of constant and alternating temperatures (25/15, 30/20, 35/25 and 40/30°C day/night). The greatest seed germination was at a constant temperature of 30°C. Seed germination was reduced at the lowest alternating temperature (25/15°C). Germination of both populations was strongly stimulated by light, suggesting a great amount of emergence of D. radulans on bare ground, such as crop seedbeds. Germination of the D. radulans population collected from a northern cotton farming system (Dalby) was more tolerant to a greater range of salt stress than the population sourced from the south (Coleambally). Seeds of both populations germinated over a wide range of pH, between 4 and 10. However, germination was the greatest in a high pH buffer solution, indicating that the species prefers to germinate in alkaline soil. These results showed that D. radulans seeds possess a wide range of tolerance mechanisms to different environmental stresses. Information obtained in this study will help in developing more sustainable and effective integrated weed management strategies for the control of this weed and weeds with similar responses in summer cropping systems, such as cotton.     

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.     

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.     

The ecological life cycle of Thlaspi perfoliatum and a comparison with published studies on Thlaspi arvense

Thlaspi perfoliatum L. is an introduced winter annual that grows in waste places, but it is not a troublesome agricultural weed. Seeds are dormant at maturity in spring, afterripen during summer and germinate in autumn. Non-dormant seeds that fail to germinate in autumn are induced into secondary dormancy by low winter temperatures and cannot germinate the following spring. These seeds afterripen during summer and germinate in autumn if conditions are suitable. Vernalization is not an absolute requirement for flowering. However, it shortens the time to flowering, and vernalized plants produce many more flowers and seeds than non-vernalized plants. Results from studies on seed germination and flowering of T. perfoliatum are compared to published studies on the life cycle of Thlaspi arvense L., an introduced species that is a serious agricultural weed which behaves both as a winter and a summer annual.     

Karrikins promote germination of physiologically dormant seeds of Chrysanthemoides monilifera ssp. monilifera (boneseed)

Physiological dormancy in weed species has significant implications for weed management, as viable seeds may persist in soil seedbanks for many years. The major stimulatory compound in smoke, karrikinolide (KAR₁), promotes germination in a range of physiologically dormant weed species allowing targeted eradication methods to be employed. Control of Chrysanthemoides monilifera ssp. monilifera (boneseed), a Weed of National Significance in Australia, may benefit from adopting such an approach. In this study, we hypothesised that seeds of C. monilifera ssp. monilifera exhibit physiological dormancy, germinate more rapidly as dormancy is alleviated, show fluctuations in sensitivity to KAR₁ and form a persistent soil seedbank. Seeds responded to 1 μM KAR₁ (40–60% germination) even during months (i.e. March, April, July, August) when seeds were observed to be more deeply dormant (control germination: 7–20%). Seeds germinated readily over a range of cooler temperatures (i.e. 10, 15, 20, 20/10 and 25/15°C) and were responsive to KAR₂ (~50% germination) as well. Eradication efforts for C. monilifera ssp. monilifera may benefit from use of karrikins to achieve synchronised germination from soil seedbanks, even at times of the year when C. monilifera ssp. monilifera seeds would be less likely to germinate, allowing more rapid depletion of the soil seedbank and targeted control of young plants.     

Temperature requirements for germination of buried seeds of Aethusa cynapium L.

Freshly-collected mature mericarps of Aethusu cynapium were dormant, but some germinated at alternating (16 h low/8 high) temperatures when the seed coverings were removed. Burial during winter increased percentage germination and the temperature range over which it took place. In late spring the range narrowed, first at low and then at higher temperatures, widening again in autumn. Moist storage at both low (4°C) and high (30°C) temperatures overcame dormancy, but exposure to 30°C inhibited subsequent germination at low temperatures. Germination of intact mericarps was consistently lower than that of de-coated seeds. The cyclic change in dormancy status of the seeds appears to interact with the restricting effects of the seed coverings and perhaps other factors in determining the consistent pattern of spring emergence in A. cynapium.     

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.     

Seed germination, seedling establishment and growth patterns of wrinklegrass (Ischaemum rugosum Salisb.)

Several laboratory and glasshouse experiments were conducted to assess seed germination, seedling establishment and growth patterns of wrinklegrass (Ischaemum rugosum Salisb.) influenced by temperature and light regimes, and chemical media. Wrinklegrass was a positively photoblastic species, and seed germination was temperature‐dependent and light‐mediated. Seeds soaked in distilled water for 24 h, or oven‐dried at the respective temperature regimes of 15, 20, 25, 30, 35, or 40°C prior to treatment in distilled water and incubated in darkness, failed to germinate. Likewise, no germination prevailed when the seeds were exposed to similar temperature regimes and treated with 0.2 m KNO₃, 5% H₂O₂ or 0.01 m HNO₃, and incubated under continuous darkness. Seeds treated with 5% H₂O₂ at 30°C, or oven‐dried and treated with 0.01% M HNO₃ at 35°C registered 10 and 20% germination. Approximately 75 and 90% of the light‐exposed seeds for all treatments germinated in the first three and six days at 25°C. No germination occurred at 15°C in the first three days after treatment. Seeds subjected to 40°C for six days after treatment recorded 36% germination. The optimum temperatures for seed germination were 25–30°C. Seed drying and soaking treatments widened the windows of the optimal temperatures for wrinklegrass germination. The acidic media of KNO₃, H₂O₂ or HNO₃ favored seed germination. Less than 5% of seed germination occurred with burial or water inundation at depths exceeding 2 cm. Seed burial or inundation at ≥2 cm depths inhibited seed germination. Seeds sown onto moist paddy soils registered ca. 50% germination. Free‐floating seeds on the water surface registered ca. 98% germination within the first six days after seeding. The mean number of seedlings that survived was inversely proportional to water depths, with close to 100% mortality at the 14 cm depths of inundation. Both plant height and seedling survival were linearly proportional to the amount of root mass of seedlings which penetrated the soil. The weed was a prolific seed producer (ca. 6000 seeds/genet or 18 000 seeds/genet per year). The vegetative and reproductive efforts of each wrinklegrass plant registered values of 0.68 and 0.32, respectively.     

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.     

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.     

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.     

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.     

Seed germination of the invasive species Piper aduncum as influenced by high temperature and water stress

Piper aduncum is a common woody weed throughout the tropics and a successful invader in many areas. As this species most often dominates roadsides and forest margins, and these habitats frequently experience extreme changes in temperature and water availability, it is important to understand how the seeds adapt to these stresses. This study investigated the effect of high temperatures and water stress, continuous, periodic or transient, on P. aduncum seed germination before or after imbibition. It was found that P. aduncum seeds exhibited only intermediate tolerance and did not germinate at temperatures above 35°C or water potentials below ?0.6 MPa. However, this species is well adapted to local conditions, with fruit ripening in the rainy season, seed desiccation tolerance to eRH 8%, rapid germination under wet conditions, higher temperature tolerance than the maximum ground temperature, germination of most seeds after 60 h continuous heat treatment or daily periodic heat treatments up to 5 h at 40°C and insensitivity to 12 h heat treatment at 40°C during the whole germination process. The intermediate tolerance to high temperature and water stress of P. aduncum, in combination with local environmental conditions in Xishuangbanna, makes P. aduncum an intermediate invader in this area.     

Ecological aspects of seed dormancy-break and germination in Heteranthera limosa (Pontederiaceae), a summer annual weed of rice fields

Summary Heteranthera limosa seeds were buried in flooded and in non-flooded soil and exposed to natural seasonal temperature changes in Lexington, Kentucky, USA. Seeds exhumed after various periods of burial ranging from 2 to 36 months were tested for germination under both flooded and non-flooded conditions. Seeds were dormant at maturity in September and became non-dormant during winter. Seeds buried in non-flooded soil during winter germinated to higher percentages and over a wider range of temperatures when tested under flooded conditions (in light) during spring and summer, than did those buried in flooded soil during winter. Thus, the water regime associated with rice culture (non-flooded in winter and flooded in summer) is optimal for dormancy-break and germination of H. limosa seeds. A portion of the buried seeds exhibited an annual dormancy/non-dormancy cycle, whereas others had a conditional dormancy/non-dormancy cycle. Regardless of the type of cycle, seeds buried in non-flooded soil retained the ability to germinate in light at high temperatures under flooded conditions throughout the summer. Thus, seeds potentially can germinate at any time during the growing season, whenever rice fields are flooded. Flooding fields during winter and/or sowing rice relatively early in the growing season may help in establishing rice before seeds of H. limosa germinate.     

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.     

Factors affecting seed germination and emergence of Aegilops tauschii

Information on seed germination and emergence ecology of Aegilops tauschii is scant, despite it being a widespread invasive weed in China. We conducted this study to determine the effects of various factors on seed germination and seedling emergence in three A. tauschii populations. Seeds germinated across a wide range of temperatures (5–35°C), with germination of over 90% at 15–20°C. Germination was completely inhibited when dry seeds were exposed to a temperature of 160°C for 5 min; a similar response was observed for pre-soaked seeds at 100°C. Light was neither required for nor inhibited germination. Germination was not significantly affected by pH. Aegilops tauschii was relatively tolerant to low osmotic potential and high salt stress: over 80% of seeds germinated at −0.3 MPa, and all three populations germinated in the presence of 400 mM salt (NaCl) although salt tolerance varied among the populations. Seeds buried at depths of 1–3 cm emerged well, but emergence was completely inhibited at depths greater than 8 cm. The addition of maize straw caused a linear reduction in seedling emergence, although the rate of reduction varied among the populations. The results of this study have contributed to understanding the requirements of A. tauschii germination and emergence and optimising an integrated management system for this weed in Huang–Huai–Hai Plain of China. In addition, our study provides data for development of models to predict the geographical distribution of this weed.