Effect of burial depth and soil water regime on the fate of Lithospermum arvense seeds in relation to burial time

Lithospermum arvense is an increasing annual weed in winter crops of the semiarid region of southern Argentina under low impact tillage systems, an agricultural practice that has become popular in recent years. Seed distribution in the soil profile under conventional tillage will change when reduced tillage is implemented, thus affecting the germination microenvironment. The effect of seed burial depth and soil water regime on field germination, enforced dormancy, innate dormancy and seed decay was studied in relation to burial time in a field experiment. In addition, the effect of burial depth on seed germination and seedling emergence was examined under laboratory controlled conditions. Field germination of buried seed ranged from 55% to 65% for shallow (2 cm) and from 5% to 30% for greater depths (20 cm). Enforced dormancy levels were significantly higher among deeper seeds. The amount of innate dormant seeds was reduced to <10% after a year of burial. Lithospermum arvense seedbanks can be classified as short-term persistent. Germination in the laboratory was unaffected by burial depth, while seedling emergence reduction was adequately described by a sigmoidal model. Results indicate that agricultural practices that accumulate L. arvense seeds near the soil surface enhance seedling recruitment.     

Seed dormancy and soil seedbank of the invasive weed Chenopodium hybridum in north‐western China

Seed dormancy and persistence in the soil seedbank play a key role in timing of germination and seedling emergence of weeds; thus, knowledge of these traits is required for effective weed management. We investigated seed dormancy and seed persistence on/in soil of Chenopodium hybridum, an annual invasive weed in north‐western China. Fresh seeds are physiologically dormant. Sulphuric acid scarification, mechanical scarification and cold stratification significantly increased germination percentages, whereas dry storage and treatments with plant growth regulators or nitrate had no effect. Dormancy was alleviated by piercing the seed coat but not the pericarp. Pre‐treatment of seeds collected in 2012 and 2013 with sulphuric acid for 30 min increased germination from 0% to 66% and 62% respectively. Effect of cold stratification on seed germination varied with soil moisture content (MC) and duration of treatment; seeds stratified in soil with 12% MC for 2 months germinated to 39%. Burial duration, burial depth and their interaction had significant effects on seed dormancy and seed viability. Dormancy in fresh seeds was released from October to February, and seeds re‐entered dormancy in April. Seed viability decreased with time for seeds on the soil surface and for those buried at a depth of 5 cm, and 39% and 10%, respectively, were viable after 22 months. Thus, C. hybridum can form at least a short‐lived persistent soil seedbank.     

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.     

Seedling emergence and seed longevity of the tree weeds Celtis sinensis and Cinnamomum camphora

Factors influencing germination and seed survival of Celtis sinensis Pers. and Cinnamomum camphora (L.) Presl. were investigated in a series of experiments conducted under natural rainfall regimes in the field and under irrigation in a growth tunnel. Germination from surface-sown seeds of C. camphora was virtually nil under natural rainfall, in contrast to almost 100% germination under irrigation. This suggests that surface-lying seeds of this species will germinate only in highly protected microsites in the field. For C. sinensis , however, in one of the two field experiments germination from surface-sown seeds was similar (≈40%) to that achieved under irrigation. Germination from intact fruits of both species was generally considerably less than from seeds. Seeds of both species were relatively short-lived under field conditions. A maximum of 1% of seeds of C. camphora remained viable at 12 months post-sowing under field conditions, whereas a maximum of 8% viable seeds of C. sinensis was detected at 24 months post-sowing in one of the field experiments.     

Seed survival and patterns of seedling emergence studies of Rottboellia cochinchinensis (Lour.) W.D. Clayton in cultivated soils

The longevity of buried Rottboellia cochinchinensis (Lour.) W.D. Clayton seed represents a major survival mechanism for the weed, enabling the persistence of a continuing source of weed seeds in crop land. The pattern of seed persistence and depletion of R. cochinchinensis in cultivated maize soils was investigated by means of (1) studies on the effect of depth and duration of burial on the viability of the weed seeds, (2) quantitative estimation of the seed population and viability in cultivated fields, and (3) the periodicity of emergence and effects of cultivation on seed germination both in the field and in the greenhouse. The results indicated that the mode of persistence was innate (8.5%) and enforced (35%) dormancy after 1 year of burial, and that the persistency component of the seed population on cultivated soils ranged from 40.60%. The weed was able to remain viable at depths of 45 cm, indicating an excellent mechanism of escaping the effects of most soil-applied herbicides, and it was shown that tillage increases the depletion rate of the weed seed reserve by 32% per year.     

Effects of the seed weight and burial depth on the seed behavior of common ragweed (Ambrosia artemisiifolia)

Common ragweed (Ambrosia artemisiifolia L.) is one of the annual plants that were described recently as invasive weeds in Europe. This species is described as an invasive plant that produces seeds that are highly variable. Its production of variably sized seeds is regarded as promoting its spread in different environments. Experiments were carried out to determine the influence of the seed weight and temperature on germination and the influence of the seed weight and burial depth on seedling emergence. The seeds were divided into a number of classes of weight and the seed weight effect on germination was evaluated by Petri dish assays. In another experiment, the seeds were buried at different depths in a clay soil/sand mix to estimate the burial effect on germination and seedling emergence. The germination level of A. artemisiifolia was high overall, between 76.8% and 94.2%. The seed germination was modified by temperature but it was not influenced by the seed weight. The amounts of germination and seedling emergence were greater for the seeds on the soil surface and decreased with an increasing burial depth, from 2 to 8 cm. No germination or emergence was observed for the seeds that were buried at 10 and 12 cm. The lightest seeds were more sensitive to burial. A greater level of seedling emergence for those seeds that were placed near the soil surface could explain the success of this species in open habitats, where the probability of deeper burial is low. After high seed production, the management of A. artemisiifolia in fields could be partly achieved through soil tillage, burying seeds below 10 cm, and not carrying out deep soil tillage the following year.     

Germination, emergence, and dormancy of Mimosa pudica

Mimosa pudica (common sensitive plant) is a problematic weed in many crops in tropical countries. Eight experiments were conducted to determine the effects of light, seed scarification, temperature, salt and osmotic stress, pH, burial depth, and rice residue on the germination, seedling emergence, and dormancy of M. pudica seeds. Scarification released the seeds from dormancy and stimulated germination, though the germination of the scarified seeds was not influenced by light. The scarification results indicate that a hard seed coat is the primary mechanism that restricts germination. The germination increased markedly with the exposure to high temperature "pretreatment" (e.g. 150°C), which was achieved by placing non-scarified seeds in an oven for 5 min followed by incubation at 35/25°C day/night temperatures for 14 days. The germination of the scarified seeds was tolerant of salt and osmotic stress, as some seeds germinated even at 250 mmol L⁻¹ NaCl (23%) and at an osmotic potential of −0.8 MPa (5%). The germination of the scarified seeds was >74% over a pH range of 5–10. The seedling emergence of the scarified seeds was 73–88% at depths of 0–2 cm and it gradually decreased with an increasing depth, with no seedling emergence at the 8 cm depth. The rice residue applied to the soil surface at rates of ≤6 t ha⁻¹ did not influence the seedling emergence and dry weight. The information gained from this study identifies some of the factors that facilitate M. pudica becoming a widespread weed in the humid tropics and might help in developing components of integrated weed management practises to control this weed.     

Germination, persistence, and emergence of flaxleaf fleabane (Conyza bonariensis [L.] Cronquist)

Flaxleaf fleabane ( Conyza bonariensis [L.] Cronquist) is a difficult-to-control weed in dryland minimum tillage farming systems in the northern grains region of Australia. Experiments under field and controlled environments were conducted to study the seed germination, emergence, and persistence of C. bonariensis . The base, optimum, and maximum temperatures for seedling germination were estimated at 4.2, 20, and 35°C, respectively, with light stimulating germination. The soil types and burial depths had significant effects on seed persistence and emergence. The persistence pattern showed an initial rapid drop, followed by a slow but steady decline over time. After 3 years of burial, there were ≈ 7.5%, 9.7%, and 1.3% viable seeds at 10, 5, and 0–2 cm soil depths, respectively. Conyza bonariensis predominantly emerged from the soil surface or from the burial depth of 0.5 cm. Very few seeds emerged from a depth of 1 cm and none emerged from ≥ 2 cm. The emergence was greater in lighter soils. On-farm monitoring of the field emergence over time in a light sandy loam soil showed that 99% emergence occurred in late autumn, early and late winter, and a small fraction of 1% emerged in mid-spring. The predominant emergence from the soil surface suggests that the fleabane problem is a result of weed species shift toward the minimum tillage systems that provide moist conditions for better emergence compared to conventional tillage systems. The seed longevity of at least 3 years in the soil and the prolonged emergence patterns between autumn and spring have posed great difficulties for the long-term management of C. bonariensis .     

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.     

Changes in germinability, dormancy and viability of Amaranthus retroflexus as affected by depth and duration of burial

Changes in dormancy and viability of Amaranthus retroflexus seeds were examined after placement in pots that were buried in the field. Seeds were placed in woven nylon envelopes on the soil surface or buried at depths of 2.5, 5 or 10 cm. After 1, 3, 6, 9 and 12 months seeds were exhumed and their germinability was tested to assess changes in dormancy and viability. Depletion of seed stocks placed on the soil surface was partly because of in situ germination that did not exceed 21% and did not vary significantly over the 12-month study period. Less germination of buried seeds occurred in situ , and seeds that did not germinate appeared to acquire dormancy. Exhumed seeds germinated readily; germinability was linearly related to the depth of burial, with those retrieved from the surface germinating least. Cyclical changes in germinability occurred during the 12-month burial period, but this influence was identical for seeds buried at all depths. Germinability was greatest after periods with the lowest mean monthly temperatures and least during the hottest periods. The stimulation of remaining ungerminated seeds exhumed at each period, by the addition of ethephon to the germination medium, provided further evidence of a seasonal acquisition of dormancy, and it was concluded that other unknown factors besides cyclical changes in seasonal temperature were responsible. Irrespective of placement, all seeds lost viability at an exponential rate over time. However, the decline was most rapid for those placed on the surface, whereas the loss in viability became less with increased depth of burial. Possible explanations for this adaptation of enhanced survival when buried are discussed.     

Changes in seed bank size and dormancy characteristics of the glyphosate-resistant biotype of goosegrass (Eleusine indica[L.] Gaertn.)

A study on changes in the seed bank size of a glyphosate ( N -(phosphonomethyl)glycine)-resistant (R) Eleusine indica biotype was carried out on a naturally occurring infestation of a young oil palm plantation. The total number of shed seeds collected was 53% of the predicted potential total number of seeds produced by the plants. Ametryn ( N -ethyl- N' -(methylethyl)-6-(methylthio)-1,3,5-triazine-2, 4-diamine) was found to be able to control the R seed population in the soil. The periodicity of seedling emergence was influenced by rainfall subsequent to the 1 month period of seed shedding. Seed burial at 2, 10 and 20 cm depths showed that 39, 33 and 79%, respectively, of viable seed persisted in the soil after 2 years. Seeds in the state of enforced dormancy played an important role in maintaining the seed population throughout the 2 year period of burial in the soil.     

Seed dynamics in populations of A vena sterilis ssp. ludoviciana

Seed populations of Avena sterilis ssp. ludoviciana (Durieu) Nyman were monitored in a naturally occurring infestation throughout its life cycle. Considering the large weed population present (298panicles m^?2), total seed production was relatively low: 3838 seeds m^?2. Only 68% of these seeds were recovered from the soil surface and a further 3% were removed with wheat grain and straw during harvest operations. The numbers of seeds from the stubble between mid-July and mid-September were relatively low (10%). Ploughing the stubble in October buried most of the recently produced seed rain and resulted in a relatively uniform vertical distribution of the seedbank. Maximum seed persistence in the soil ranged from 27 to 43 months (depending on the experimental technique used to do the study). Seed decline followed an exponential pattern on a yearly basis, with the greatest decline taking place between October and April (57–90% in year 1 and 10–40% in year 2), Between May and September the buried seed populations remained practically constant. Seedbank depletion was primarily due to seedling production (25%) and ‘lethal’ germination (24%). Although the depth of burial had very little effect on seed survival, the mode of seed disappearance was closely related to their depth in the soil. Seed depletion through ‘lethal’ germination increased with increasing depth in the soil, whereas depletion through seedling emergence decreased with increasing depth.     

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.     

Longevity of seeds of four annual grass and two dicotyledon weed species as related to placement in the soil and straw disposal technique

Persistence of seeds of the four winter annual grass species, Alopecurus myosuroides , Bromus sterilis , Bromus hordeaceus and Vulpia myuros , were studied under field conditions. The studies also included the two dicotyledon species, Tripleurospermum inodorum and Galium aparine . Seed samples were stored in the field at different depths in the plough layer for a year. Following germination, seedling emergence was taken as an indicator of field persistence. In a supplementary investigation, seed samples were placed on the soil surface or buried at 2 cm for a month and then tested for viability. Persistence of the two Bromus species was very short, irrespective of depth and duration of burial. Persistence of the other species was positively correlated with incorporation of seeds to a few centimetres depth. Increasing the depth of incorporation below 2 cm had a variable influence on persistence. Persistence of seeds placed directly on the soil surface was short, whether they were left there for a year or a month. Whether seeds at the soil surface were left uncovered or covered with chopped straw had limited influence on seed persistence. However, straw cover significantly increased persistence of A. myosuroides seeds and there was the same tendency with T. inodorum .     

Annual cycles of germinability and differences between primary and secondary dormancy in buried seeds of Echinochloa crus-galli

The seasonal changes in percentage of dormant seeds of Echinochloa crus-galli in the field were recorded for 4 years. The lots of seeds were wrapped in nylon fabric, buried 20 cm under the grass sward and exhumed at monthly intervals. The proportion of seeds germinating under light conditions at a constant temperature of 25 °C fluctuated between 0% and 96%, with maxima in May–July and minima in September–November. Small between-year differences in the course of summer dormancy induction and its winter termination were probably caused by variation of weather conditions.
Attributes of dormancy innate to seeds after maturation (primary dormancy) and dormancy induced in buried seeds during the summer (secondary dormancy) were compared by investigating the rate of dormancy termination during storage of (a) dry seeds at 25 °C, (b) imbibed seeds at 5°C and (c) in seeds buried under field conditions during October–June. Percentage of germination increased faster in secondary than primary dormant seeds at both constant 25 °C and 5 °C. The seeds with primary and secondary dormancy also differed in the response to `germination pre-treatment', a 10-day exposure of imbibed seeds at 25 °C that causes germination of the non-dormant fraction of seed materials. After this treatment the time to resuming germination in primary dormant seeds was substantially increased, whereas the secondary dormant seeds were much less affected. Annual variation in the proportion of germinable seeds explains the low efficiency of autumn soil cultivation for decreasing reserves of E. crus-galli seeds in the soil.     

Germination characteristics and emergence time of annual Bromus species of differing weediness in Sweden

The germination ecology of four annual Bromus species, which differ in weediness on arable land in southern Sweden, was investigated. The most problematic species is Bromus sterilis , while Bromus hordeaceus frequently occurs on arable land. In contrast, Bromus arvensis is a rare weed, and Bromus tectorum is found infrequently in fields despite being a widespread ruderal species. Five experiments were conducted to identify germination characteristics that could explain differences in habitat and abundance: (i) intraspecific variation in dormancy level; (ii) germination response to different light conditions; (iii) light and temperature interactions at germination; (iv) timing of seedling emergence; and (v) seed persistence in soil. Bromus sterilis and B. tectorum behaved similarly in all tests. For both these species, there were large differences in dormancy level among populations and strong inhibition of germination by light. In addition, emergence from seeds sown on the soil surface was both delayed and reduced compared with buried seeds. In contrast, B. hordeaceus and B. arvensis showed generally weak dormancy, and germination was only slightly inhibited by light. It was concluded that germination characteristics alone do not explain the differences in weediness between these four species.     

Germination ecology of Leptochloa chinensis: a new weed in the Italian rice agro-environment

Leptochloa chinensis is a new weed that has been found with increasing frequency in Italian rice paddies. The germination ecology of L. chinensis seeds was studied in order to investigate the development mechanisms and survival strategy of this weed in rice paddies of northern Italy. Leptochloa chinensis seeds showed no dormancy and exhibited germination even in anoxic conditions. Germination was strongly influenced by temperature (minimum around 15°C; optimal 25–35°C) and light (phytochrome dependent). Temperature fluctuation caused an increase of seed germination in the dark. Seed burial also strongly inhibited germination and emergence of this species. At 5 cm seed burial only 5% of seedlings emerged in flooded conditions, while at the same depth, but with no flooding, no seedling emergence was observed. This phenomenon was not due to oxygen depletion, as germination was not inhibited by complete anoxia, as demonstrated by the fact that some seedlings did emerge in flooding conditions when water was no deeper than 6 cm. Seed burial and concomitant flooding induced an unusual germination: first coleoptile emergence and subsequently emergence of the radicle was observed. The possible exploitation of this knowledge for weed management is discussed.     

Comparison of seed germination of four Artemisia species (Asteraceae) in northeastern Inner Mongolia, China

Two psammophile-dominated Artemisia semi-shrubs (A.wudanica and A.halodendron) and two annual Artemisia forbs (A.sieversiana and A.scoparia) bear significant ecological functions in Horqin Sandy Land,but systematical information on their achenes' germination is very limited.A set of studies were conducted to evaluate seed germination responses to storage periods and methods,different temperatures,lights conditions and sand burial depths,in order to determine inter-specific germination variation in the same genus and to explain how the species adapt to its microhabitat.Fresh achenes of A.wudanica,A.halodendron and A.sieversiana showed high germination capacities,but those of A.scoparia had obvious innate dormancy,which could be broken by chilling and dry storage,especially long-term dry storage.Achene germination of the two semi-shrubs preferred lower temperature fluctuation (10 to 22oC) and was not sensitive to light.But the two annuals preferred higher temperature fluctuation (34 to 22oC) and strong light for their achene germination.These four Artemisia species showed similar responses to sand burial,i.e.soil surface was most favorable for seedling emergence,and the deeper the sand burial,the fewer the seedling emergence.For the two semi-shrubs,their microhabitats are sand dunes with high temperature and intense light,which are not favorable for germination and seedling survival.Only rainfall contributes to temporary decrease of temperature and then triggers germination.We deduced that germination is not the main but a supplementary reproductive mode for the two semi-shrubs in sand dunes.For the two annuals,achene germination is the only reproductive mode,but different responses have been developed for microhabitat adaptation.For A.sieversiana,high germination capacities in wide temperature ranges and all light conditions could improve its competition and advancement in the wettest microhabitats.For A.scoparia,obvious innate dormancy of fresh achenes and germination inhabitation under unfavorable conditions are important adaptation to environmental disturbances.     

Seed biology of Mikania micrantha in Viti Levu,Fiji

Mikania micrantha (mile‐a‐minute) reproduces both by seed and vegetatively. A study to determine the possible pollinators, seed production rates, temperature and salinity limitations to germination and its seedbank size and persistence was conducted in Viti Levu, Fiji. Representatives of the Diptera had the greatest percentage of all floral visits (38%), followed by Hymenoptera (34%) and Lepidoptera (27%), while the honeybee was the most recurring visitor (18% of all visits). Flower heads (capitula) within the inflorescence commonly formed four viable seeds, resulting in 60 820 filled seed being produced per m². However, the seedbanks formed were not massive (600 seed m^?2) and they were moderately persistent (T₅₀: 1–3 years). Seed germination from both high and moderate rainfall regions occurred rapidly, under a wide range of temperature regimes, with no primary dormancy being observed. This study indicates that the seed reproductive success of M. micrantha in the two rainfall regions of Fiji is due to a number of factors, including the production of large numbers of flowers, successful pollination by local insects and the subsequent production of a large number of viable seeds. These seeds have high viability, no dormancy and are capable of forming small‐to‐medium seedbanks that are moderately long‐lived. In addition, seeds can germinate under a wide range of temperature and salinity conditions. This knowledge on seed production, biology, longevity and salt tolerance is vital in the development of management plans of M. micrantha in Fiji.     

Burial depth and cultivation influence emergence and persistence of Phalaris paradoxa seed in an Australian sub-tropical environment

Emergence and persistence characteristics of Phalaris paradoxa seeds in no- and minimum-till situations and at different burial depths were studied in a sub-tropical environment. Three experiments were carried out using naturally shed seeds. In the first experiment, seedlings emerged from May through to September each year, although the majority of seedlings emerged in July. In the second experiment with greater seed density, cultivation in March of each year stimulated seedling emergence, altered the periodicity of emergence and accelerated the decline of seeds in the seedbank compared with plots that received no cultivation. The majority of seedlings in the cultivated plots emerged in May whereas the majority of seedlings in the undisturbed plots emerged in July. Emergence accounted for only 4–19% of the seedbank in both experiments over 2 years. Seed persistence was short in both field experiments, with less than 1% remaining 2 years after seed shed. In the third experiment, burial depth and soil disturbance significantly influenced seedling emergence and persistence of seed. Seedlings emerged most from seed mixed in the top 10 cm when subjected to annual soil disturbance, and from seed buried at 2.5 and 5.0 cm depths in undisturbed soil. Emergence was least from seed on the soil surface, and buried at 10 and 15 cm depths in undisturbed soil. Seeds persisted longest when shed onto the soil surface and persisted least when the soil was tilled. These results suggest that strategic cultivation may be a useful management tool, as it will alter the periodicity of emergence allowing use of more effective control options and will deplete the soil seedbank more rapidly.