Priming rice seeds (soaking followed by drying) or soaking just before sowing improved emergence from flooded soil, reduced membrane damage from ROS and hastened carbohydrate mobilization. Most benefit was to lines with a superior ability to germinate in flooded soil even when untreated.
Background and aims
Early flooding helps control weeds but reduces seedling establishment in direct-seeded rice (Oryza sativa). When combined with appropriate management practices, the use of genotypes that better tolerate flooding during emergence can enhance crop establishment in flood-prone areas. Management options include seed pre-treatment and we tested the influence of pre-soaking for 24 h prior to sowing or of priming (soaking for 24 or 48 h followed by drying).
The effects on seedling establishment after 21-day flooding of pre-soaking seeds for 24 h before sowing and/or of priming seeds were examined together with physiological responses connected with reactive oxygen scavenging. Seeds of four lines with contrasting abilities to tolerate flooding at the germination stage were compared. Seeds were primed using KCl solutions (48 h) or water (24 h) and pre-soaked using water. Lipid peroxidation and activities of reactive oxygen-scavenging enzymes were measured in seeds before sowing. Carbohydrate mobilization in germinating seeds and seedling growth were also monitored at intervals.
Seed pre-treatment by pre-soaking or by priming increased survival of flooding and accelerated and improved seedling establishment, especially in tolerant genotypes. Primed seeds had less lipid peroxidation and higher superoxide dismutase (SOD) and catalase (CAT) activities than non-primed seeds. Amylase activities and starch breakdown were also hastened in primed seeds. Survival after flooding was positively correlated with amylase activity but negatively correlated with the extent of lipid peroxidation.
Pre-soaking and priming improved seedling establishment in flooded soil, enhanced the capacity to scavenge reactive oxygen species in seeds by increasing SOD and CAT activities, and hastened carbohydrate mobilization. Tolerant genotypes responded better to these treatments, emphasizing the effectiveness of combining genetic tolerance with appropriate seed pre-treatment to improve seedling establishment of rice sown in flooded soils.
Background and aims
In recent years, Cyperus rotundus has become a problem weed in lowland rice (Oryza sativa) grown in rotation with vegetables in the Philippines. As the growth of C. rotundus is commonly suppressed by prolonged flooding, the ability of the weed to grow vigorously in flooded as well as upland conditions suggests that adapted ecotypes occur in these rotations. Studies were conducted to elucidate the mechanisms that permit C. rotundus to tolerate flooded soil conditions.
Upland and lowland ecotypes of C. rotundus were compared in terms of growth habit, carbohydrate reserves and metabolism, and activities of enzymes involved in alcoholic fermentation – alcohol dehydrogenase (ADH) and pyruvate decarboxylase (PDC).
The lowland ecotype has much larger tubers than the upland ecotype. Prior to germination, the amylase activity and total non-structural carbohydrate content in the form of soluble sugars were greater in the tubers of lowland plants than in those of upland C. rotundus. At 24 h after germination in hypoxic conditions, PDC and ADH activities in the lowland plants increased, before decreasing at 48 h following germination. In contrast, ADH and PDC activities in the upland plants increased from 24 to 48 h after germination.
Tolerance of lowland C. rotundus of flooding may be attributed to large carbohydrate content and amylase activity, and the ability to maintain high levels of soluble sugars in the tubers during germination and early growth. This is coupled with the modulation of ADH and PDC activities during germination, possibly to control the use of carbohydrate reserves and sustain substrate supply in order to avoid starvation and death of seedlings with prolonged flooding.
Anoxia; ethanol fermentation; flooding tolerance; nutsedge; Cyperus rotundus; Pasteur effect; weed ecology
Comparing a lowland and an upland ecotype of Cyperus rotundus, the former had greater carbohydrate reserves in tubers, thicker roots and stems with larger air spaces and, under hypoxia, it maintained relatively lower activities of alcohol dehydrogenase and lactate dehydrogenase.
Background and aims
Purple nutsedge (Cyperus rotundus L.) is a major weed of upland crops and vegetables. Recently, a flood-tolerant ecotype evolved as a serious weed in lowland rice. This study attempted to establish the putative growth and physiological features that led to this shift in adaptation.
Tubers of upland C. rotundus (ULCR) and lowland C. rotundus (LLCR) ecotypes were collected from their native habitats and maintained under the respective growth conditions in a greenhouse. Five experiments were conducted to assess the variation between the two ecotypes in germination, growth and tuber morphology when grown in their native or ‘switched’ conditions. Carbohydrate storage and mobilization, and variation in anaerobic respiration under hypoxia were compared.
Tubers of LLCR were larger than those of ULCR, with higher carbohydrate content, and larger tubers developed with increasing floodwater depth. Stems of LLCR had larger diameter and proportionally larger air spaces than those of ULCR: a method of aerating submerged plant parts. The LLCR ecotype can also mobilize and use carbohydrate reserves under hypoxia, and it maintained relatively lower and steadier activity of alcohol dehydrogenase (ADH) as a measure of sustained anaerobic respiration. In contrast, ADH activity in ULCR increased faster upon a shift to hypoxia and then sharply decreased, suggesting depletion of available soluble sugar substrates. The LLCR ecotype also maintained lower lactate dehydrogenase activity under flooded conditions, which could reduce chances of cellular acidosis.
These adaptive traits in the LLCR ecotype were expressed constitutively, but some of them, such as tuber growth and aerenchyma development, are enhanced with stress severity. The LLCR ecotype attained numerous adaptive traits that could have evolved as a consequence of natural evolution or repeated management practices, and alternative strategies are necessary because flooding is no longer a feasible management option.
Background and Aims
Flooding slows seed germination, imposes fatalities and delays seedling establishment in direct-seeded rice. This study describes responses of contrasting rice genotypes subjected to flooding or low oxygen stress during germination and discusses the basis of tolerance shown by certain cultivars.
In one set of experiments, dry seeds were sown in soil and either watered normally or flooded with 10 cm of water. Seedling survival and shoot and root growth were assessed and seed portions of germinating seedlings were assayed for soluble sugars and starch concentrations. The whole germinating seedlings were assayed for amylase and peroxidase activities and for ethylene production. Activities of enzymes associated with anaerobic respiration were examined and gene expression was analysed separately with seeds germinating under different amounts of dissolved oxygen in dilute agar.
Flooding during germination reduced survival but to a lesser extent in tolerant genotypes. Starch concentration in germinating seeds decreased while sugar concentration increased under flooding, but more so in tolerant genotypes. Amylase activity correlated positively with elongation (r = 0·85 for shoot and 0·83 for root length) and with plant survival (r = 0·92). Tolerant genotypes had higher amylase activity and higher RAmy3D gene expression. Ethylene was not detected in seeds within 2 d after sowing, but increased thereafter, with a greater increase in tolerant genotypes starting 3 d after sowing. Peroxidase activity was higher in germinating seeds of sensitive genotypes and correlated negatively with survival.
Under low oxygen stress, tolerant genotypes germinate, grow faster and more seedlings survive. They maintain their ability to use stored starch reserves through higher amylase activity and anaerobic respiration, have higher rates of ethylene production and lower peroxidase activity as germinating seeds and as seedlings. Relevance of these traits to tolerance of flooding during germination and early growth is discussed.
Amylase; anoxia; crop establishment; direct-seeded rice; ethylene; flooding; germination; hypoxia; Oryza sativa
Rice is semi-aquatic, adapted to a wide range of hydrologies, from aerobic soils in uplands to anaerobic and flooded fields in waterlogged lowlands, to even deeply submerged soils in flood-prone areas. Considerable diversity is present in native rice landraces selected by farmers over centuries. Our understanding of the adaptive features of these landraces to native ecosystems has improved considerably over the recent past. In some cases, major genes associated with tolerance have been cloned, such as SUB1A that confers tolerance of complete submergence and SNORKEL genes that control plant elongation to escape deepwater. Modern rice varieties are sensitive to flooding during germination and early growth, a problem commonly encountered in rainfed areas, but few landraces capable of germination under these conditions have recently been identified, enabling research into tolerance mechanisms. Major QTLs were also identified, and are being targeted for molecular breeding and for cloning. Nevertheless, limited progress has been made in identifying regulatory processes for traits that are unique to tolerant genotypes, including faster germination and coleoptile elongation, formation of roots and leaves under hypoxia, ability to catabolize starch into simple sugars for subsequent use in glycolysis and fermentative pathways to generate energy. Here we discuss the state of knowledge on the role of the PDC-ALDH-ACS bypass and the ALDH enzyme as the likely candidates effective in tolerant rice genotypes. Potential involvement of factors such as cytoplasmic pH regulation, phytohormones, reactive oxygen species scavenging and other metabolites is also discussed. Further characterization of contrasting genotypes would help in elucidating the genetic and biochemical regulatory and signaling mechanisms associated with tolerance. This could facilitate breeding rice varieties suitable for direct seeding systems and guide efforts for improving waterlogging tolerance in other crops.
anaerobic germination; alcoholic fermentation; ALDH; pyruvate dehydrogenase bypass; hypoxia; direct seeding; flooding; submergence tolerance
Poor seed germination and early seedling growth associated with urea-induced soil ammonia volatilization are major constraints in the adoption of dry direct-seeded rice. To directly examine soil ammonia volatilization and its damage to seed germination and early seedling growth of dry direct-seeded rice when urea is applied at seeding, two Petri-dish incubation experiments and a field experiment were conducted. Ammonia volatilization due to urea application significantly reduced seed germination and early seedling growth of dry direct-seedling rice. NBPT significantly reduced ammonia volatilization following urea application. The application of ammonium sulfate, instead of urea at seeding, may mitigate poor crop establishment of dry direct-seeded rice. Root growth of dry direct-seeded rice was more seriously inhibited by soil ammonia volatilization than that of shoot. Results suggest that roots are more sensitive to soil ammonia toxicity than shoots in dry direct-seeded rice system when N is applied as urea at seeding.
Background and Aims
Soil phosphorus (P) solubility declines sharply when a flooded soil drains, and an important component of rice (Oryza sativa) adaptation to rainfed lowland environments is the ability to absorb and utilize P under such conditions. The aim of this study was to test the hypothesis that rice cultivars differ in their P responses between water regimes because P uptake mechanisms differ.
Six lowland rice cultivars (three considered tolerant of low P soils, three sensitive) were grown in a factorial experiment with three water regimes (flooded, moist and flooded-then-moist) and four soil P levels, and growth and P uptake were measured. Small volumes of soil were used to maximize inter-root competition and uptake per unit root surface. The results were compared with the predictions of a model allowing for the effects of water regime on P solubility and diffusion.
The plants were P stressed but not water stressed in all the water regimes at all P levels except the higher P additions in the flooded soil. The cultivar rankings scarcely differed between the water regimes and P additions. In all the treatments, the soil P concentrations required to explain the measured uptake were several times the concentration of freely available P in the soil.
The cultivar rankings were driven more by differences in growth habit than specific P uptake mechanisms, so the hypothesis cannot be corroborated with these data. Evidently all the plants could tap sparingly soluble forms of P by releasing a solubilizing agent or producing a greater root length than measured, or both. However, any cultivar differences in this were not apparent in greater net P uptake, possibly because the restricted rooting volume meant that additional P uptake could not be converted into new root growth to explore new soil volumes.
Oryza sativa; rainfed lowland; phosphorus efficiency; root morphology; solubilization; rice cultivar
We review strategies of sexual and asexual reproduction and persistence in plants of flood-prone Central Amazonia. Adaptations in response to the strong instability of these environments are highlighted together with the importance of river connectivity for species dispersal and persistence.
The Central Amazonian floodplain forests are subjected to extended periods of flooding and to flooding amplitudes of 10 m or more. The predictability, the length of the flood pulse, the abrupt transition in the environmental conditions along topographic gradients on the banks of major rivers in Central Amazonia, and the powerful water and sediment dynamics impose a strong selective pressure on plant reproduction systems.
In this review, we examine how the hydrological cycle influences the strategies of sexual and asexual reproduction in herbaceous and woody plants. These are of fundamental importance for the completion of the life cycle. Possible constraints to seed germination, seedling establishment and formation of seed banks are also covered. Likewise, we also discuss the importance of river connectivity for species propagation and persistence in floodplains.
The propagation and establishment strategies employed by the highly diversified assortment of different plant life forms result in contrasting successional stages and a zonation of plant assemblages along the flood-level gradient, whose species composition and successional status are continuously changing not only temporally but also spatially along the river channel.
Background and Aims
Reducing damage to rice seedlings caused by flash flooding will improve the productivity of rainfed lowland rice in West Africa. Accordingly, the morphological and physiological responses of different forms of rice to complete submergence were examined in field and pot experiments to identify primary causes of damage.
To characterize the physiological responses, seedlings from a wide genetic base including Oryza sativa, O. glaberrima and interspecific hybrids were compared using principle component analysis.
Important factors linked to flash-flood tolerance included minimal shoot elongation underwater, increase in dry matter weight during submergence and post-submergence resistance to lodging. In particular, fast shoot elongation during submergence negatively affected plant growth after de-submergence. Also shoot-elongating cultivars showed a strong negative correlation between dry matter weight of the leaves that developed before submergence and leaves developing during submergence.
Enhancement of shoot elongation during submergence in water that is too deep to permit re-emergence by small seedlings represents a futile escape strategy that takes place at the expense of existing dry matter in circumstances where underwater photosynthetic carbon fixation is negligible. Consequently, it compromises survival or recovery growth once flood water levels recede and plants are re-exposed to the aerial environment. Tolerance is greater in cultivars where acceleration of elongation caused by submergence is minimal.
Africa; flash floods; Oryza glaberrima; rainfed lowland; rice; shoot elongation; stress tolerance; submergence
Evolved herbicide resistance (EHR) is an important agronomic problem and consequently a food security problem, as it jeopardizes herbicide effectiveness and increases the difficulty and cost of weed management. EHR in weeds was first reported in 1970 and the number of cases has accelerated dramatically over the last two decades. Despite 40 years of research on EHR, why some weeds evolve resistance and others do not is poorly understood. Here we ask whether weed species that have EHR are different from weeds in general. Comparing taxonomic and life history traits of weeds with EHR to a control group (“the world's worst weeds”), we found weeds with EHR significantly over-represented in certain plant families and having certain life history biases. In particular, resistance is overrepresented in Amaranthaceae, Brassicaceae and Poaceae relative to all weeds, and annuality is ca. 1.5 times as frequent in weeds with EHR as in the control group. Also, for perennial EHR weeds, vegetative reproduction is only 60% as frequent as in the control group. We found the same trends for subsets of weeds with EHR to acetolactate synthase (ALS), photosystem II (PSII), and 5-enolpyruvylshikimate-3-phosphate (EPSP) synthase-inhibitor herbicides and with multiple resistance. As herbicide resistant crops (transgenic or not) are increasingly deployed in developing countries, the problems of EHR could increase in those countries as it has in the USA if the selecting herbicides are heavily applied and appropriate management strategies are not employed. Given our analysis, we make some predictions about additional species that might evolve resistance.
It has been suggested that genetically modified herbicide-tolerant crops may benefit biodiversity because spraying of crops may be delayed until later in the growing season, allowing weeds to grow during the early part of the year. This provides an enhanced resource for arthropods, and potentially benefits birds that feed on these. Thus, this technology could enhance biodiversity. Using a review of weed phenologies and a population model, we show that many weeds are unlikely to benefit because spraying is generally delayed insufficiently late in the season to allow most to set seed. The positive effects on biodiversity observed in trials lasting one or two seasons are thus likely to be transient. For one weed of particular significance (Chenopodium album, fat hen) we show that it is unlikely that the positive effects observed could be maintained by inputs of seed during other parts of the rotation. However, we find preliminary evidence that if spraying can be ceased earlier in the season, then a viable population of late-emerging weeds could be maintained. This strategy could benefit weeds in both genetically modified (GM) and non-GM crops, but would probably lead to reduced inputs in GM systems compared with conventional ones.
Plant transgenic technology has been widely utilized for engineering crops for trait improvements and for production of high value proteins such as pharmaceuticals. However, the unintended spreading of commercial transgenic crops by pollination and seed dispersal is a major concern for environmental and food safety. Simple and reliable containment strategies for transgenes are highly desirable. Here we report a novel method for creating selectively terminable transgenic rice. In this method, the gene(s) of interest is tagged with a RNA interference cassette, which specifically suppresses the expression of the bentazon detoxification enzyme CYP81A6 and thus renders transgenic rice to be sensitive to bentazon, a herbicide used for rice weed control. We generated transgenic rice plants by this method using a new glyphosate resistant 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) gene from Pesudomonas putida as the gene of interest, and demonstrated that these transgenic rice plants were highly sensitive to bentazon but tolerant to glyphosate, which is exactly the opposite of conventional rice. Field trial of these transgenic rice plants further confirmed that they can be selectively killed at 100% by one spray of bentazon at a regular dose used for conventional rice weed control. Furthermore, we found that the terminable transgenic rice created in this study shows no difference in growth, development and yield compared to its non-transgenic control. Therefore, this method of creating transgenic rice constitutes a novel strategy of transgene containment, which appears simple, reliable and inexpensive for implementation.
The UK Farm Scale Evaluations (FSEs) have shown that the use of broad spectrum herbicides on genetically modified herbicide-tolerant (GMHT) crops can have dramatic effects on weed seed production compared to management of conventional varieties. Here, we use FSE data and information on bird diets to determine how GMHT cropping might change the food resources available to farmland birds. More than 60 fields of each of four crops, spring- and winter-sown oilseed rape, beet and maize, were split, one half being sown with a conventional variety, the other with a GMHT variety. Seed rain from weeds known to be important in the diets of 17 granivorous farmland bird species was measured under the two treatments. In beet and spring oilseed rape, rain of weed seeds important in the diets of 16 bird species was significantly reduced in GMHT compared to conventional halves; for no species did it increase. In winter oilseed rape, rain of weed seeds important in the diets of 10 species was significantly reduced in GMHT halves; for only one species did it increase significantly. By contrast, in maize, rain of weed seeds important in the diets of seven species was significantly greater in GMHT halves; for no species was it reduced. Treatment effects for the total weed seed energy available to each bird species were very similar to those for seed rain alone. Measuring the effects on individual bird species was outside the scope of this study. Despite this, these results suggest that should beet, spring and winter rape crops in the UK be largely replaced by GMHT varieties and managed as in the FSEs, this would markedly reduce important food resources for farmland birds, many of which declined during the last quarter of the twentieth century. By contrast, GMHT maize would be beneficial to farmland birds.
genetically modified herbicide-tolerant crops; weed seeds; farmland birds; bird food
Tropospheric ozone is one of the major drivers of global change. This stress factor alters plant growth and development. Ozone could act as a selection pressure on species communities composition, but also on population genetic background, thus affecting life history traits. Our objective was to evaluate the consequences of prolonged ozone exposure of a weed community on phenotypic traits of Spergulaarvensis linked to persistence. Specifically, we predicted that the selection pressure exerted by high ozone concentrations as well as the concomitant changes in the weed community would drive population adaptive changes which will be reflected on seed germination, dormancy and longevity. In order to test seed viability and dormancy level, we conducted germination experiments for which we used seeds produced by S. arvensis plants grown within a weed community exposed to three ozone treatments during four years (0, 90 and 120 ppb). We also performed a soil seed bank experiment to test seed longevity with seeds coming from both the four-year ozone exposure experiment and from a short-term treatment conducted at ambient and added ozone concentrations. We found that prolonged ozone exposure produced changes in seed germination, dormancy and longevity, resulting in three S. arvensis populations. Seeds from the 90 ppb ozone selection treatment had the highest level of germination when stored at 75% RH and 25 °C and then scarified. These seeds showed the lowest dormancy level when being subjected to 5 ºC/5% RH and 25 ºC/75% followed by 5% RH storage conditions. Furthermore, ozone exposure increased seed persistence in the soil through a maternal effect. Given that tropospheric ozone is an important pollutant in rural areas, changes in seed traits due to ozone exposure could increase weed persistence in fields, thus affecting weed-crop interactions, which could ultimately reduce crop production.
Background and Aims
In the Amazonian floodplains plants withstand annual periods of flooding which can last 7 months. Under these conditions seedlings remain submerged in the dark for long periods since light penetration in the water is limited. Himatanthus sucuuba is a tree species found in the ‘várzea’ (VZ) floodplains and adjacent non-flooded ‘terra-firme’ (TF) forests. Biochemical traits which enhance flood tolerance and colonization success of H. sucuuba in periodically flooded environments were investigated.
Storage carbohydrates of seeds of VZ and TF populations were extracted and analysed by HPAEC/PAD. Starch was analysed by enzyme (glucoamylase) degradation followed by quantification of glucose oxidase. Carbohydrate composition of roots of VZ and TF seedlings was studied after experimental exposure to a 15-d period of submersion in light versus darkness.
The endosperm contains a large proportion of the seed reserves, raffinose being the main non-structural carbohydrate. Around 93 % of the cell wall storage polysaccharides (percentage dry weight basis) in the endosperm of VZ seeds was composed of mannose, while soluble sugars accounted for 2·5%. In contrast, 74 % of the endosperm in TF seeds was composed of galactomannans, while 22 % of the endosperm was soluble sugars. This suggested a larger carbohydrate allocation to germination in TF populations whereas VZ populations allocate comparatively more to carbohydrates mobilized during seedling development. The concentration of root non-structural carbohydrates in non-flooded seedlings strongly decreased after a 15-d period of darkness, whereas flooded seedlings were less affected. These effects were more pronounced in TF seedlings, which showed significantly lower root non-structural carbohydrate concentrations.
There seem to be metabolic adjustments in VZ but not TF seedlings that lead to adaptation to the combined stresses of darkness and flooding. This seems to be important for the survival of the species in these contrasting environments, leading these populations to different directions during evolution.
Carbohydrate reserves; Himatanthus sucuuba; submergence tolerance; floodplains; galactomannans; raffinose; population differentiation; Amazon; storage
A new oxyvinylglycine has been identified as a naturally occurring herbicide that irreversibly arrests germination of the seeds of grassy weeds, such as annual bluegrass (Poa annua), without significantly affecting the growth of established grass seedlings and mature plants or germination of the seeds of broadleaf plant species (dicots). Previously, Pseudomonas fluorescens WH6 and over twenty other rhizosphere bacteria were isolated and selected for their ability to arrest germination of P. annua seeds. The Germination-Arrest Factor (GAF, 1) responsible for this developmentally specific herbicidal action has now been isolated from the culture filtrate of P. fluorescens WH6. Purification of this highly polar, low molecular weight natural product allowed its structure to be assigned as 4-formylaminooxy-l-vinylglycine based on NMR spectroscopic and mass spectrometric data, in combination with d/l-amino acid oxidase reactions to establish the absolute configuration. Assay results for P. annua inhibition by related compounds known to regulate plant growth are presented, and a cellular target for 1 is proposed. Furthermore, using bioassays, TLC, and capillary NMR spectroscopy, it has been shown that GAF (1) is secreted by all other herbicidally-active rhizosphere bacteria in our collection.
Echinochloaoryzicola(syn.E. phyllopogon) is an exotic weed of California rice paddies that has evolved resistance to multiple herbicides. Elimination of seedlingsthroughcertain weed control methods can limit the spread of this weed, but is contingent on accurate predictions of germination and emergence timing, which are influenced by seed dormancy levels.In summer annuals, dormancy can often be relieved through stratification, a period of prolonged exposure to cold and moist conditions.We used population-based threshold models to quantify the effects of stratification on seed germination of four E. Oryzicola populations at a range of water potential (Ψ) and oxygen levels. We also determined how stratification temperatures, moisture levels and durations contributed to dormancy release. Stratification released dormancy by decreasing base Ψ and hydrotimerequired for germination and by eliminating any germination sensitivity to oxygen. Stratification also increased average germination rates (GR), which were used as a proxy for relative dormancy levels. Alternating temperatures nearly doubled GR in all populations, indicating that seeds could be partially dormant despite achieving high final germination percentages. Stratification at Ψ = 0 MPa increased GR compared to stratification at lower water potentials, demonstrating that Ψ contributed to regulating dormancy release. Maximum GR occurred after 2-4 weeks of stratification at 0 MPa; GR were often more rapid for herbicide-resistant than for herbicide-susceptible seeds, implying greater dormancy in the latter. Manipulation of field conditions to promote dormancy alleviation of E. oryzicola seeds might improve the rate and uniformity of germination for seed bank depletion through seedling weed control. Our results suggest field soil saturation in winter would contribute towards E. oryzicola dormancy release and decrease the time to seedling emergence.
The Farm Scale Evaluations (FSEs) showed that genetically modified herbicide-tolerant (GMHT) cropping systems could influence farmland biodiversity because of their effects on weed biomass and seed production. Recently published results for winter oilseed rape showed that a switch to GMHT crops significantly affected weed seedbanks for at least 2 years after the crops were sown, potentially causing longer-term effects on other taxa. Here, we seek evidence for similar medium-term effects on weed seedbanks following spring-sown GMHT crops, using newly available data from the FSEs.
Weed seedbanks following GMHT maize were significantly higher than following conventional varieties for both the first and second years, while by contrast, seedbanks following GMHT spring oilseed rape were significantly lower over this period. Seedbanks following GMHT beet were smaller than following conventional crops in the first year after the crops had been sown, but this difference was much reduced by the second year for reasons that are not clear. These new data provide important empirical evidence for longer-term effects of GMHT cropping on farmland biodiversity.
Farm Scale Evaluations; arable weeds; farmland biodiversity
Higher plants are aerobic organisms which suffer from the oxygen deficiency imposed by partial or total submergence. However, some plant species have developed strategies to avoid or withstand severe oxygen shortage and, in some cases, the complete absence of oxygen (tissue anoxia) for considerable periods of time.
Rice (Oryza sativa) is one of the few plant species that can tolerate prolonged soil flooding or complete submergence thanks to an array of adaptive mechanisms. These include an ability to elongate submerged shoot organs at faster than normal rates and to develop aerenchyma, allowing the efficient internal transport of oxygen from the re-emerged elongated shoot to submerged parts. However, rice seeds are able to germinate anaerobically by means of coleoptile elongation. This cannot be explained in terms of oxygen transport through an emerged shoot. This review provides an overview of anoxic rice germination that is mediated through coleoptile rather than root emergence.
Although there is still much to learn about the biochemical and molecular basis of anaerobic rice germination, the ability of rice to maintain an active fermentative metabolism (i.e. by fuelling the glycolytic pathway with readily fermentable carbohydrates) is certainly crucial. The results obtained through microarray-based transcript profiling confirm most of the previous evidence based on single-gene studies and biochemical analysis, and highlight new aspects of the molecular response of the rice coleoptile to anoxia.
Anoxia; coleoptile; fermentative metabolism; germination; hypoxia; Oryza sativa; rice
Currently, the UK has no procedure for the approval of novel agricultural
practices that is based on environmental risk management principles. Here, we
make a first application of the ‘bow-tie’ risk management
approach in agriculture, for assessment of land use changes, in a case study of
the introduction of genetically modified herbicide tolerant (GMHT) sugar beet.
There are agronomic and economic benefits, but indirect environmental harm from
increased weed control is a hazard. The Farm Scale Evaluation (FSE) trials
demonstrated reduced broad-leaved weed biomass and seed production at the field
scale. The simplest mitigation measure is to leave a proportion of rows
unsprayed in each GMHT crop field. Our calculations, based on FSE data, show
that a maximum of 2% of field area left unsprayed is required to mitigate weed
seed production and 4% to mitigate weed biomass production. Tilled margin
effects could simply be mitigated by increasing the margin width from 0.5 to
1.5 m. Such changes are cheap and simple to implement in farming
practices. This case study demonstrates the usefulness of the bow-tie risk
management approach and the transparency with which hazards can be addressed. If
adopted generally, it would help to enable agriculture to adopt new practices
with due environmental precaution.
genetically modified crops; environmental risk assessment; sugar beet; mitigation; biodiversity
We compared the seedbanks, seed rains, plant densities and biomasses of weeds under two contrasting systems of management in beet, maize and spring oilseed rape. Weed seedbank and plant density were measured at the same locations in two subsequent seasons. About 60 fields were sown with each crop. Each field was split, one half being sown with a conventional variety managed according to the farmer's normal practice, the other half being sown with a genetically modified herbicide-tolerant (GMHT) variety, with weeds controlled by a broad-spectrum herbicide. In beet and rape, plant densities shortly after sowing were higher in the GMHT treatment. Following weed control in conventional beet, plant densities were approximately one-fifth of those in GMHT beet. In both beet and rape, this effect was reversed after the first application of broad-spectrum herbicide, so that late-season plant densities were lower in the GMHT treatments. Biomass and seed rain in GMHT crops were between one-third and one-sixth of those in conventional treatments. The effects of differing weed-seed returns in these two crops persisted in the seedbank: densities following the GMHT treatment were about 20% lower than those following the conventional treatment. The effect of growing maize was quite different. Weed density was higher throughout the season in the GMHT treatment. Late-season biomass was 82% higher and seed rain was 87% higher than in the conventional treatment. The difference was not subsequently detectable in the seedbank because the total seed return was low after both treatments. In all three crops, weed diversity was little affected by the treatment, except for transient effects immediately following herbicide application.
We evaluated the effects of the herbicide management associated with genetically modified herbicide-tolerant (GMHT) winter oilseed rape (WOSR) on weed and invertebrate abundance and diversity by testing the null hypothesis that there is no difference between the effects of herbicide management of GMHT WOSR and that of comparable conventional varieties. For total weeds there were few treatment differences between GMHT and conventional cropping, but large and opposite treatment effects were observed for dicots and monocots. In the GMHT treatment, there were fewer dicots and more monocots than in conventional crops. At harvest, dicot biomass and seed rain in the GMHT treatment were one-third of that in the conventional, while monocot biomass was threefold greater and monocot seed rain almost fivefold greater in the GMHT treatment than in the conventional. These differential effects persisted into the following two years of the rotation. Bees and butterflies that forage and select for dicot weeds were less abundant in GMHT WOSR management in July. Year totals for Collembola were greater under GMHT management. There were few other treatment effects on invertebrates, despite the marked effects of herbicide management on the weeds.
genetically modified crops; biodiversity; oilseed rape; canola; herbicide management
In recognition of the 200th anniversary of Charles Darwin's birth, this short article on flooding stress acknowledges not only Darwin's great contribution to the concept of evolution but also to the study of plant physiology. In modern biology, Darwin-inspired reductionist physiology continues to shed light on mechanisms that confer competitive advantage in many varied and challenging environments, including those where flooding is prevalent.
Mild flooding is experienced by most land plants but as its severity increases, fewer species are able to grow and survive. At the extreme, a highly exclusive aquatic lifestyle appears to have evolved numerous times over the past 120 million years. Although only 1–2% of angiosperms are aquatics, some of their adaptive characteristics are also seen in those adopting an amphibious lifestyle where flooding is less frequent. Lowland rice, the staple cereal for much of tropical Asia falls into this category. But, even amongst dry-land dwellers, or certain of their sub-populations, modest tolerance to occasional flooding is to be found, for example in wheat. The collection of papers summarized in this article describes advances to the understanding of mechanisms that explain flooding tolerance in aquatic, amphibious and dry-land plants. Work to develop more tolerant crops or manage flood-prone environments more effectively is also included. The experimental approaches range from molecular analyses, through biochemistry and metabolomics to whole-plant physiology, plant breeding and ecology.
Abiotic stress; adaptation; anoxia; Charles Darwin; environmental stress; evolution; flooding; hypoxia; rice; submergence; wetlands
The host suitability to Ditylenchus destructor of seven common weed species in peanut (Arachis hypogaea) fields in South Africa was determined. Based on the number of nematodes per root unit, white goosefoot (Chenopodium album), feathertop chloris (Chloris virgata), purple nutsedge (Cyperus rotundus), jimson weed (Datura stramonium), goose grass (Eleusine indica), khaki weed (Tagetes minuta), and cocklebur (Xanthium strumarium) were poor hosts. Ditylenchus destructor survived on all weed species; population densities increased in peanut hulls and caused severe damage to seeds of peanut grown after weeds. Roots of purple nutsedge left in the soil suppressed populations of D. destructor and root and pod development in peanut grown after the weed. However, nematode populations in peanut hulls and seeds were not suppressed. Some weed species, especially purple nutsedge which is common in peanut fields, can be used to indicate the presence of D. destructor in the absence of peanut.
Arachis hypogaea; Chenopodium album; Chloris virgata; cocklebur; Cyperus rotundus; Datura stramonium; Ditylenchus destructor; Eleusine indica; feathertop chloris; goose grass; host status; jimson weed; khaki weed; peanut; purple nutsedge; South Africa; Tagetes minuta; Xanthium strumarium
Background and Aims
Seed survival in the soil contributes to population persistence and community diversity, creating a need for reliable measures of soil seed bank persistence. Several methods estimate soil seed bank persistence, most of which count seedlings emerging from soil samples. Seasonality, depth distribution and presence (or absence) in vegetation are then used to classify a species' soil seed bank into persistent or transient, often synthesized into a longevity index. This study aims to determine if counts of seedlings from soil samples yield reliable seed bank persistence estimates and if this is correlated to seed production.
Seeds of 38 annual weeds taken from arable fields were buried in the field and their viability tested by germination and tetrazolium tests at 6 month intervals for 2·5 years. This direct measure of soil seed survival was compared with indirect estimates from the literature, which use seedling emergence from soil samples to determine seed bank persistence. Published databases were used to explore the generality of the influence of reproductive capacity on seed bank persistence estimates from seedling emergence data.
There was no relationship between a species' soil seed survival in the burial experiment and its seed bank persistence estimate from published data using seedling emergence from soil samples. The analysis of complementary data from published databases revealed that while seed bank persistence estimates based on seedling emergence from soil samples are generally correlated with seed production, estimates of seed banks from burial experiments are not.
The results can be explained in terms of the seed size–seed number trade-off, which suggests that the higher number of smaller seeds is compensated after germination. Soil seed bank persistence estimates correlated to seed production are therefore not useful for studies on population persistence or community diversity. Confusion of soil seed survival and seed production can be avoided by separate use of soil seed abundance and experimental soil seed survival.
Arable weeds; Bifora testiculata; Carthamus lanatus; Centaurea solstitialis; longevity index; seed bank persistence; soil seed bank