Induction of extreme light sensitivity in buried weed seeds and its role in the perception of soil cultivations

Abstract. Light, probably acting through the photo-receptor phytochrome, promotes germination of weed seeds when the soil is disturbed by tillage operations. A short period of burial is shown to induce an enormous ∼10000-fold increase in light sensitivity in the seeds of the arable weed Datura ferox which is interpreted as a natural transition to the 'very-low-fluence' mode of phytochrome action. Field experiments indicated that germination of buried seeds may be triggered by millisecond-exposures to sunlight and suggested a key role for the process of sensitization in the mechanisms whereby light requiring seeds detect the occurrence of soil cultivation events in arable lands.     

Modelling field emergence patterns in arable weeds

A model was developed to simulate weed emergence patterns after soil cultivation. In the model, the consecutive processes of dormancy release, germination and pre-emergence growth were modelled in separate modules. Input variables of the model were: date of soil cultivation, soil temperature and soil penetration resistance. Output variables of the model were: seedling density and timing of seedling emergence. The model was parameterized for Polygonum persicaria , Chenopodium album and Spergula arvensis with data from previous field and laboratory experiments. The model was evaluated with data from an experiment, in which emergence of P. persicaria , C. album and S. arvensis was monitored in field plots that were cultivated once only, at one of five dates in the spring. At the same time as the field observations on seedling emergence, seasonal changes in seed dormancy of the buried weed seeds were assessed by testing the germination of seed lots that were buried in envelopes. From a comparison between field observations and simulated data, it appeared that the model overestimated the rate of dormancy release in spring, whereas germination and pre-emergence growth were simulated well. In general, therefore, both the numbers of emerging seedlings and the timing of emergence could be predicted accurately, when dormancy was not simulated but introduced from experimental data. Improvement of predictions of field emergence of weeds should mainly focus on increasing the precision of the simulation of dormancy release. Close correlations were found between seedbed temperature and both the extent and rate of seedling emergence, but analysis with the simulation model revealed that they were only partly based on causal relationships, so that they have limited predictive value.     

Seed Germination Ecology of Feather Lovegrass [Eragrostis tenella (L.) Beauv. Ex Roemer & J.A. Schultes]

Feather lovegrass [Eragrostis tenella (L.) Beauv. Ex Roemer & J.A. Schultes] is a C₄ grass weed that has the ability to grow in both lowland and upland conditions. Experiments were conducted in the laboratory and screenhouse to evaluate the effect of environmental factors on germination, emergence, and growth of this weed species. Germination in the light/dark regime was higher at alternating day/night temperatures of 30/20 °C (98%) than at 35/25 °C (83%) or 25/15 °C (62%). Germination was completely inhibited by darkness. The osmotic potential and sodium chloride concentrations required for 50% inhibition of maximum germination were -0.7 MPa and 76 mM, respectively. The highest seedling emergence (69%) was observed from the seeds sown on the soil surface and no seedlings emerged from seeds buried at depths of 0.5 cm or more. The use of residue as mulches significantly reduced the emergence and biomass of feather lovegrass seedlings. A residue amount of 0.5 t ha^-1 was needed to suppress 50% of the maximum seedlings. Because germination was strongly stimulated by light and seedling emergence was the highest for the seeds sown on the soil surface, feather lovegrass is likely to become a problematic weed in zero-till systems. The knowledge gained from this study could help in developing effective and sustainable weed management strategies.     

The fate of Datura ferox seeds in the soil as affected by cultivation, depth of burial and degree of maturity

Seeds of Datura ferox were collected in soybean fields, grouped into four categories according to the degree of fruit maturity and placed just level with the soil surface or buried to a depth of 7 or 15 cm. Seed survival after 8 months was c. 30% when the seeds were left on the soil surface, but between 40 and 90% when the seeds were buried. The riper seeds were the more persistent. A similar pattern was observed after 20 months. Seedling emergence was negatively related with the degree of seed maturity and depth of burial, but it was never great enough to explain seed losses. In another experiment seeds from ripe capsules were superficially sown and the soil was: (0) left undisturbed, (1) cultivated in late winter or (2) cultivated in late winter and late spring. In all cases there was a small flush of seedlings at the beginning of the first spring; other flushes occurred only after soil cultivation and were larger during the second spring than during the first. In the plots cultivated twice a year (2) the seedlings that emerged in the first spring represented c. 4% of the initial seed bank. Survival after 20 months was about 25% in all treatments. In an arable field under soybean cropping the seedling flushes during the spring months were related to the pattern of soil cultivation. The seedlings that emerged after crop drilling represented c. 18% of the seed bank (estimated before drilling). Implications for weed management are discussed.     

SOME EXPERIMENTS AND FIELD OBSERVATIONS ON THE GERMINATION OF WILD OAT (AVENA FATUA AND A. LUDOVICIANA) SEEDS IN SOIL AND THE EMERGENCE OF SEEDLINGS

The distinguishing characters of Avena fatua and A. ludoviciana are described.
Pot experiments and field observations showed that most seeds of A. fatua germinated in spring and a few in autumn; hardly any germinated in summer or winter. Seeds of A. ludoviciana germinated in winter only. The greatest depth of sowing from which seedlings of either species reached the surface was 9 in., but seedlings from this depth were weak and yellow when they first appeared. A. ludoviciana gave more and sturdier seedlings than A. fatua from 6 and 9 in. There was no evidence of induced dormancy in seeds of A. fatua buried at depths down to 20 in. Germination of this species was hastened by monthly cultivation of the soil. The maximum survival of A. fatua was 3 years in pots and slightly longer in the field; seeds of A. ludoviciana in pots survived only 2 years. The two or three seeds of each spikelet of A. ludoviciana germinated in turn, starting with the largest, but the interval between germination of successive seeds varied.
Seedlings from freshly sown seeds of both species were more vigorous than seedlings from seeds which had been buried for a year or more. Germination and subsequent growth of both species took place in soil of pH 4.5 to 7.0 approx.     

Stimulation of weed seed germination by 1–(3-chlorophthalirnido) cyclohexanecarboxamide (AC 94377)

The N-substituted phthalimide, AC 94377, promoted the germination of dry-stored seeds of 17 out of 24 weed species in Petri-dish tests maintained at 22°C. In a further test it was able to substitute for light and/or alternating temperatures in promoting the germination of some species. AC 94377 was more active than GA, in stimulating Solanum nigrum seed to germinate. When mixed with soil in the laboratory, AC 94377 again promoted germination of S. nigrum but the level and persistence of activity differed between soils. In the glasshouse, with soil containing a natural weed seed population, more seedlings emerged from treated than from untreated soil but this number represented fewer than half of the apparently viable seeds present. Field applications of AC 94377 promoted the germination of hand-sown seeds of certain weed species but did not increase the number of seedlings emerging from the natural weed seed bank.     

Crop and weed emergence patterns in relation to time of cultivation and rainfall

In experiments over 3 years, separate field plots were cultivated once only at 2-wk intervals. Cultivation resulted in a flush of weed seedlings, representing usually less than 6% of the apparently viable seeds in the top 10 cm of soil and with a species composition which varied with the time of year. Subsequent flushes coincided in timing and extent with those on soil which had been undisturbed and were related to significant rainfall events. In each year there were periods when lack of soil moisture restricted emergence. At these times, seeds were released from dormancy by cultivation but germination of some or all of them was delayed. There was a general correspondence in the emergence patterns of weeds and of carrot and onion sown after each cultivation. On occasions when the seedbed was drying out, establishment of the comparatively rapidly-germinating radish and cabbage was greater than that of carrot, onion or weeds. Effective initiation of germination of both crops and weeds was traceable to rainfall events. Emergence was reduced or delayed on more than half of the 45 cultivation occasions during the three growing seasons, and the results emphasise the importance of soil moisture in determining the extent and timing of seedling emergence.     

不同光照处理下青藏高原克隆植物黄帚橐吾(Ligulariavirgaurea)种子大小对其幼苗生长的影响

以高寒草甸克隆植物黄帚橐吾为实验材料,通过遮荫网模拟植被遮荫,研究种子大小与萌发及幼苗生长能力的关系和幼苗对光照条件的反应。结果表明：（1）在自然光照下,黄帚橐吾种子大小对种子萌发的影响显著,大种子的萌发率高于小种子。遮荫生境下,大、小种子萌发率有所降低,但遮荫对小种子萌发的影响比大种子显著。小种子的萌发率下降了近1/8,而大种子的萌发率仅下降了1/11。（2）黄帚橐吾种子大小对幼苗生物量积累影响显著,大种子幼苗总生物量（TB）大于小种子幼苗的。但生物量的分配与播种时间相关,播种后60 d,在自然光照条件下,大种子幼苗对根生物量的分配大于小种子幼苗,而对叶生物量的分配则正好相反。在遮荫环境中,大、小种子幼苗普遍对根的生物量分配增加,大种子幼苗根冠比（R/S）大于小种子幼苗。（3）黄帚橐吾种子大小对幼苗的生长也有明显影响。在自然光照下,小种子幼苗的相对生长速率（RGR）较大于大种子幼苗,但叶面积比率（LAR）、叶面积干质量比（SLA）、叶干质量（LWR）差别不明显。在遮荫条件下,幼苗的LAR、SLA、LWR显著增加,但大、小种子幼苗间差异不显著,幼苗的RGR减小,小种子幼苗的减小趋势大于大种子幼苗。     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

Effect of light environment during soil disturbance on germination and emergence pattern of weeds

This paper describes results from experiments which investigated the effects of light intensity during soil disturbance on germination and emergence pattern of weeds. Different emergence patterns were demonstrated for seeds which are instantly flash induced compared to seeds which are induced to germinate by integrating a weak light signal over a period of time. A reduced and delayed emergence is achieved after a disturbance in darkness compared to a soil disturbance in daylight. The increased emergence after soil disturbance in daylight is due to additional plants originating from seeds placed at a soil depth in the pots where daylight cannot penetrate and induce seeds to germinate, but which are induced during the short exposure period. A close relationship between soil disturbance intensity and number of weed plants emerging was found in field experiments with shallow harrowings. It was also shown that a portion of the increased number of seedlings arising when soil disturbance is carried out in daylight, compared to soil cultivation in darkness, originates from seeds germinating from deeper soil layers, resulting in a deeper average germination depth.     

紫茎泽兰土壤种子库特征及其对幼苗的影响

研究紫茎泽兰(Eupatorium adenophorum)土壤种子库特征及其对紫茎泽兰种子的萌发、幼苗命运的影响, 在综合治理紫茎泽兰入侵危害及防止紫茎泽兰的继续扩散等方面有着重要的意义。该文在攀枝花紫茎泽兰入侵严重的地区, 通过采集果园、放牧灌丛以及禁牧灌丛3种不同生境紫茎泽兰土壤种子库的样本, 以说明不同干扰程度下紫茎泽兰种子在土壤中的分布状况。通过野外调查并结合盆栽实验, 初步研究了紫茎泽兰土壤种子库基本特征以及光照和种子在土壤中的埋藏深度等对紫茎泽兰幼苗的影响。结果表明: 1)果园、放牧灌丛和禁牧灌丛等3种干扰程度不同生境的深层种子量占总种子量的比例分别为56.44%、46.96%和24.86% (p=0.006), 这说明土壤深层种子量大小与干扰成正比, 干扰越大, 深层次紫茎泽兰种子量占总种子量的比重越大。2)播种在0、1和5 cm深度的种子萌发率分别为64.67%、22.67%和13.33%, 即种子埋藏越深, 萌发率越低, 不同层次种子萌发率差异极显著(p=0.00); 幼苗死亡率分别为27.95%、0和0, 表层种子萌发的幼苗有较高的死亡率, 而由埋藏在深层的种子萌发的幼苗没有死亡, 土壤表层发芽的幼苗与不同埋藏深度种子萌发的幼苗之间死亡率差异极显著(p=0.00)。3)在无遮蔽、半遮蔽和全遮蔽3种不同情况下, 紫茎泽兰幼苗的死亡率分别为72.15%、30.38%和4.87%, 定居率分别为6.66%、33.99%和46.67%, 即遮蔽程度越高, 死亡率越低, 定居率越高, 不同处理之间死亡率和定居率差异均极显著(p=0.00)。研究结果暗示, 强光可能是导致紫茎泽兰幼苗死亡的重要原因, 人类活动的干扰可能导致更多的紫茎泽兰种子进入土壤深层, 从而改变了紫茎泽兰土壤种子库的结构。由于土壤深层种子比表层种子具有更强的抵抗强光照射等不良环境因子影响的能力, 所萌发的幼苗成活率高, 表明其具有更高的繁殖效率, 因此人类活动干扰是紫茎泽兰入侵后难以根除的原因之一。     

Survival and seed to seedling transmission of Alternaria linicola on linseed

Alternaria linicola survived as thick-walled chlamydospores in hyphal or conidial cells on infected linseed stem debris, either on the soil surface or buried in the soil, during the period between the harvest of linseed and sowing of the following crop (September - April). Conidia produced on these debris under favourable conditions were not only viable but also pathogenic to young linseed seedlings. Infected stem debris increased the incidence of infected seedlings which emerged from infected seed (incidence of A. linicola 1% to 28%), especially if the debris was on the soil surface rather than buried. A. linicola also survived between successive linseed crops on infected volunteer linseed plants which survived the low temperatures in winter and on the weed Veronica persica. A. linicola was more effectively transmitted from infected seeds to seedlings at temperatures 15–25°C than at 10°C. The incidence of the disease on seedlings which emerged from infected seed was positively correlated with the amount of seed-borne inoculum, whereas the proportion of seedlings which emerged was negatively correlated with the incidence of A. linicola on the seed.     

The influence of straw disposal and cultivation regime on the population dynamics ofBromus sterilis

The effects of straw-burning and cultivation regime upon the population dynamics of a naturally-occurring infestation of Bromus sterilis were investigated over a period of 11 months. Both grain and straw of the original crop were seriously contaminated with seeds of this species. An initial population of 12 635 seeds m^-2 declined by 85% between July and late August in uncultivated stubble without straw-burning, but only 44% of seeds gave rise to seedlings. A further 10% had produced seedlings by late December and another 5% emerged between February and April. By April no viable seed remained. Straw-burning destroyed 97% of the ungerminated seeds on the soil surface and reduced seedling numbers by 94%. However, those seedlings which did survive formed many tillers and produced considerable quantities of seed. Shallow cultivation reduced the seed population by 34% and in April there were 47% fewer seedlings on these plots than on direct drilled ones. Ploughing to a depth of 20 cm eradicated the weed; and although buried seeds germinated, they failed to emerge. Seedlings emerging in spring failed to flower before harvest. Detailed laboratory investigation showed that although B. sterilis does not have a pronounced requirement for vernalisation, chilling did accelerate flowering, while long-day photoperiods were required for panicle extension. Only germinated seeds responded to the vernalisation stimulus.     

杂草种子寿命的研究

1976—1978年先后把稗、看麦娘、藜、猪殃殃等24种农田主要杂草种子分别埋入水田和旱田(棉麦)的土壤内,每年挖出一部分种子进行发芽试验,测定其存活率,8年试验结果表明,经4年后稗草种子在水田内还有34.2—52.5%发芽率,至第7年全部死亡;而在旱田内经2年,只有8.7%的发芽率,第三年死亡率达100.0%;看麦娘、硬草、棒头草种子在水田内2年发芽率分别为39.3%,62.7%、33.3%,而在旱田内2年后死亡率达100.0%;猪殃殃、波斯婆婆纳的种子在旱田中2年内的发芽率为57.0%、80.0%,但在水田内其死亡率达98.7%、100.0%;马齿苋的种子在旱田内经7年发芽率为33.3%;而水田内为23.0%。在水、旱不同条件下,不同杂草种子的寿命是不一样的,这将为水旱轮作防除农田杂草提供了理论依据。     

Seed-transmission in the ecology of nematode-borne viruses

Virus-free populations of vector nematodes can acquire tomato black ring (TBRV), raspberry ringspot (RRV) and arabis mosaic (AMV) viruses from weed seedlings grown from virus-carrying seed. When soils from fields where nematode-borne viruses occurred naturally were air-dried to kill vector nematodes and then moistened, TBRV and RRV occurred commonly in the weed seedlings that grew, but AMV occurred only rarely. Similar tests did not detect tobacco ringspot, grapevine fanleaf or tobacco rattle viruses in weed seeds in the single soil studied in each instance, although these three viruses are also seed-borne in some of their hosts. Many weed species, when infected experimentally, readily transmit TBRV and RRV to their seed, but the viruses were much commoner in naturally occurring seed of some of these species than of others. These discrepancies between the frequency of seed-transmission of viruses from experimentally infected plants and the extent of natural occurrence of infected seed seem largely to reflect the host preferences of the vectors. Infective Longidorus elongatus kept in fallow soil retained TBRV and RRV only up to 9 weeks. When weed seeds in the soil were then allowed to germinate, the nematodes reacquired virus from the infected seedlings. Some weed species were better than others as sources of virus. Persistence of these viruses in fields through periods of fallow or fasting of the vector therefore depends on a continuing supply of infected seedlings produced by virus-containing weed seeds. This is probably less true of viruses like AMV and grapevine fanleaf, which persist for 8 months or more in their vectors (Xiphinema spp.). A few seeds containing TBRV and RRV were found in soils free of vector nematodes, suggesting that the viruses are disseminated in weed seed. This probably explains how TBRV and RRV have reached a large proportion of L. elongatus populations in eastern Scotland.     

FIELD GERMINATION AND SEEDLING GROWTH OF CH AND CL PROGENY OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The germination behavior and early growth of chasmogamous (CH) and cleistogamous (CL) progeny of Impatiens capensis were investigated in two eastern Nebraska stands. Field germination of families of buried seeds was scored in mid-April. In one stand a significantly higher proportion of CL seeds germinated than CH seeds while no significant differences were observed in the second stand. Among-family variation in germination rates was significant in both stands. The time course of field germination was similar for CL and CH seeds. A sample of the germinated CL and CH seedlings, matched in size, was planted individually in pots and grown in the greenhouse for five wk. Early growth of CL and CH seedlings was similar. In addition, CL seedlings were as variable as CH seedlings for four morphological traits. Overall, a large fitness advantage for CH progeny was not detected in the germination and early growth of Impatiens capensis seedlings.     

Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Fire and the transition to seedlings

Fire, through soil heating effects, causes flushes of seed germination in Acacia suaveolens. Optimal temperatures for germination are between 60 and 80°C for any duration, or up to 100°C for durations less than 1 h. Exposure to temperatures less than 60°C leaves seeds dormant and viable, whilst seed death occurs in increasing proportions with increasing exposure to temperatures greater than 80°C. A field study of temperatures in the soil under simulated burns showed that the innate seed dormancy in A. suaveolens would only be broken for seeds up to a depth of 1 cm in ‘cool’ or 4 cm in ‘hot’ burns. In the hot burns some of the seeds in the top 1 cm of the soil were killed by excessive heating. These simulated burns most resemble cool and moderate/high intensity wildfires, respectively. Seeds can emerge from depths up to 8 cm and, for any seeds buried deeper than this, the probability of emergence is progressively reduced down to nil at 14 cm. Seeds buried between 5 and 10 cm will be heated sufficiently to break their dormancy only in a very high intensity wildfire. Seeds buried between 5 and 10 cm deep mostly occur in nests of an ant, Pheidole sp. Field observations of emergent seedlings confirm that post-fire emergence is concentrated over a small range of soil depths directly related to the intensity and duration of heating that occurs, whilst occasional seedlings may appear from greater or lesser depths largely dependent upon the spatial heterogeneity of soil heating in natural fires.     

Phytochrome-mediated long-term memory of seeds

The question is how long phytochrome, stored within the cytoplasm of plant diaspores, may stimulate their germination. This question arose from the observation that soil cultivations in darkness for weed control gave inconsistent results. Namely, after a single nighttime or daytime cultivation during spring and summer, differences in weed emergence became hardly detectable after a period of six weeks. However, after nighttime and daytime cultivations in late autumn, emergence differences persisted for up to nine months. To examine whether this differing memory effect is phytochrome-mediated, seeds of Chenopodium album and Stellaria media were sown in pots with wet peat, either in daylight or after sunset. In the latter, seeds were irradiated with far-red light for one day prior to being covered and buried. For more than two years the far-red irradiated seeds produced significantly reduced emergence, indicating that germination and emergence of weeds in the field may be supported by maternal far-red absorbing seed phytochrome B(fr) over several months or even years. This conclusion allows refining of the strategy of lightless tillage.     

Germination and emergence characteristics of triazine-susceptible and triazine-resistant biotypes of Solanum nigrum

1. Seedling emergence patterns of triazine-susceptible and triazine-resistant Solanum nigrum in the field were studied in Wageningen, the Netherlands. Emergence patterns were similar in the first year, but in the second year resistant seedlings emerged faster and the number of resistant seedlings was higher. To explain emergence patterns, a germination experiment was carried out.
2. Seeds from two populations with triazine-susceptible and -resistant biotypes were buried in late autumn and exhumed monthly during spring. Germination was assessed in incubators at different constant temperatures.
3. The lowest temperatures for germination of seeds from the Achterberg population ranged from 20°C on 1 February to 10°C on 1 May for the susceptible biotype, and from 15°C on 1 February to 10°C on 1 May for the resistant biotype. The lowest temperatures for germination of seeds from the Zelhem population ranged from 25°C on 1 February to 10°C on 1 May for the susceptible biotype, and from 15°C on 1 February to 10°C on 1 May for the resistant biotype. The minimum germination temperature of seeds from the resistant biotype appeared to be lower than that of the susceptible biotype.
4. Emergence patterns in the field could be explained by soil temperature and different minimum germination temperature requirements of seeds from the triazine-susceptible and -resistant biotype. This knowledge can be used to manage triazine-resistant biotypes of S. nigrum by the timing of soil cultivation.