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Herbicides are the most commonly used means of controlling weeds. Recently, there has been growing concern over the potential impacts of global climate change, specifically, increasing temperatures and elevated carbon dioxide (CO 2 ) concentrations, on the sensitivity of weeds to herbicides. Here, glyphosate response of both Conyza canadensis and Chenopodium album was evaluated under different environmental conditions. Reduced glyphosate sensitivity was observed in both species in response to increased temperature, elevated CO 2 level, and the combination of both factors. Increased temperature had greater effect on plant survival than elevated CO 2 level. In combination, high temperature and elevated CO 2 level resulted in loss of apical dominance and rapid necrosis in glyphosate-treated plants. To investigate the mechanistic basis of reduced glyphosate sensitivity, translocation was examined using 14 C-glyphosate. In plants that were subjected to high temperatures and elevated CO 2 level, glyphosate was more rapidly translocated out of the treated leaf to shoot meristems and roots than in plants grown under control conditions. These results suggest that altered glyphosate translocation and tissue-specific sequestration may be the basis of reduced plant sensitivity. Therefore, overreliance on glyphosate for weed control under changing climatic conditions may result in more weed control failures.

Chenopodium album and C. murale are cosmopolitan, annual weed species of notable economic importance. Their unique biological features, including high reproductive capacity, seed dormancy, high persistence in the soil seed bank, the ability to germinate and grow under a wide range of environmental conditions and abiotic stress tolerance, help these species to infest diverse cropping systems. C. album and C. murale grow tall and absorb nutrients very efficiently. Both these species are allelopathic in nature and, thus, suppress the germination and growth of native vegetation and/or crop plants. These weed species infest many agronomic and horticultural crops and may cause > 90% loss in crop yields. C. album is more problematic than C. murale as the former is more widespread and infests more number of crops, and it also acts as an alternate host of several crop pests. Different cultural and mechanical methods have been used to control these weed species with varying degrees of success depending upon the cropping systems and weed infestation levels. Similarly, allelopathy and biological control have also shown some potential, especially in controlling C. album. Several herbicides have been successfully used to control these species, but the evolution of wide-scale herbicide resistance in C. album has limited the efficacy of chemical control. However, the use of alternative herbicides in rotation and the integration of chemicals and biologically based control methods may provide a sustainable control of C. album and C. murale.

Background: Herbicide hormesis may play a role in the evolution of weed resistance by increasing resistance selection. A standard herbicide rate may be subtoxic to resistant plants and make them more fit than untreated plants. If this increase in fitness is ultimately expressed in reproductive traits, resistance genes can accumulate more rapidly and exacerbate resistance evolution by magnifying the selection differential between resistant and sensitive plants. The hypothesis of hormetically enhanced reproductive fitness was studied for a photosystem II (PSII) target-site resistant (TSR) biotype of Chenopodium album exposed to the triazinone metamitron in comparison with its wild-type., Results: Both biotypes showed an initial hormetic growth increase at different doses leading to fitness enhancements of between 19% and 61% above untreated plants. However, hormetic effects only resulted in higher fitness at maturity in resistant plants with a maximum stimulation in seed yield of 45% above untreated plants. Applying realistic metamitron rates, reproductive fitness of resistant plants was increased by 15-32%., Conclusions: Agronomically relevant doses of metamitron induced considerable hormesis in a PSII-TSR C. album genotype leading to enhanced relative fitness through reproductive maturity. This increase in relative fitness suggests an impact on resistance selection and can compensate for the oft-reported fitness costs of the mutation studied. Field rates of herbicides can, thus, not only select for resistant plants, but also enhance their reproductive fitness. The finding that herbicide hormesis can be eco-evolutionary important may have important implications for understanding the evolution of herbicide resistance in weeds. © 2018 Society of Chemical Industry., (© 2018 Society of Chemical Industry.)

Salt stress is one of the most dramatic abiotic stresses that induce oxidative and osmotic stress simultaneously. Salt stress is known to be more effective in reducing growth and yield of glycophytes; however, halophytes are able to withstand salt stress. Nonetheless, variability exists among different halophytic plants species from different plant families. Chenopodium album belongs to Chenopodiacea family and is known as weed in many regions of world; however, it is a very interesting halophytic plant. Little research has conducted so far by considering C. album as model plant to study salt stress tolerance mechanisms. This article attempts to compile current literature in order to explain C. album salt stress tolerance mechanism and to highlight the knowledge gap relating to salt stress tolerance mechanism in C. album. Briefly, C. album has remarkable ability of seed dimorphism, sodium exclusion, and potassium retention. C. album further tolerates salt stress by increasing redox potential associated with high production of osmolytes and antioxidants.

Chenopodium album has a non-photosynthetic chlorophyll protein known as the water-soluble chlorophyll (Chl)-binding protein (WSCP). The C. album WSCP (CaWSCP) is able to photoconvert the chlorin skeleton of Chl a into a bacteriochlorin-like skeleton. Reducing reagents such as β-mercaptoethanol or dithiothreitol inhibit photoconversion, indicating that S-S bridge(s) in CaWSCP are quite important for it. Recently, we found that the mature region of CaWSCP contains five cysteine residues; Cys2, Cys30, Cys48, Cys63, and Cys144. To identify which cysteine residues are involved in the photoconversion, we generated five mutants (C2S, C30S, C48S, C63S, and C144S) by site-directed mutagenesis. Interestingly, C48S, C63S, and C144S mutants showed the same Chl-binding activity and photoconvertibility as those of the recombinant wild-type CaWSCP-His, while the C2S and C30S mutants completely lost Chl-binding activity. Our findings indicated that the S-S bridge between Cys2 and Cys30 in each CaWSCP subunit is essential for Chl-binding activity.

Production of chloroplast-localized small heat-shock proteins (Cp-sHSP) is correlated with increased thermotolerance in plants. Ecotypic variation in function and expression of Cp-sHSPs was analyzed in two Chenopodium album ecotypes from cool vs. warm-temperate USA habitats [New York (NY) and Mississippi (MS) respectively]. P(et) was more heat tolerant in the MS than the NY ecotype, and MS ecotype derived proportionally greater protection of P(et) by Cp-sHSP during high temperatures. Four genes encoding Cp-sHSPs were isolated and characterized: CaHSP25.99n (NY-1) and CaHSP26.23n (NY-2) from NY ecotype, and CaHSP26.04m (MS-1) and CaHSP26.26m (MS-2) from MS ecotype. The genes were nearly identical in predicted amino-acid sequence and hydrophobicity. Gene expression analysis indicated that MS-1 and MS-2 transcripts were constitutively expressed at low levels at 25 °C, while no NY-1 and NY-2 transcripts were detected at this temperature. Maximum accumulation of NY-1 and NY-2 transcripts occurred at 33 °C and 40 °C for MS-1 and MS-2. Immunoblot analysis revealed that (1) protein expression was highest at 37 °C in both ecotypes, but was greater in MS than NY ecotype at 40 °C; and (2) import of Cp-sHSP into chloroplasts was more heat-labile in NY ecotype. The higher expression of one isoform in MS ecotype may contribute to its enhanced thermotolerance. Absence of correlation between protein and transcript levels, suggests the post-transcriptional regulation is occurring. Promoter analysis of these genes revealed significant variations in heat-shock elements (HSE), core motifs required for heat-shock-factor binding. We propose a correlation between unique promoter architecture, Cp-sHSP expression and thermotolerance in both ecotypes., (Published by Elsevier Masson SAS.)

As soil salinity constitutes a major threat to agriculture in the world, to cope with this problem, much emphasis has been focused on the response mechanism of halophytes or salt-tolerant species to salinity stress. In the present study, C. album was treated with long-term NaCl and KCl stress, then several parameters were assayed in leaves as follows: inductively coupled plasma atomic emission spectrometry (ICP-AES) was employed to measure the K, Na content; semi-quantitative RT-PCR was used to investigate the expression level of three genes which were related to ion transport on the vacuolar membrane-NHX, VP1 and VAP-C. In addition, the mechanism for the effect of different salinity on K, Na content was preliminarily discussed. The results were as follows: (1) Under lower concentration of NaCl stress, C. album had a preferential uptake of potassium and exclusion of sodium, and thus maintained a low concentration of sodium in cells of the leaf; (2) Under higher concentration of NaCl (300 mmolx L(-1)), C. album was able to tolerate excessive amounts of sodium in cells and kept the higher K/Na ratio in cytoplasm by compartmentalizing Na ions into the vacuole via ion transporter system located on vacuole membrane, (3) Much K ions and total ions (including sodium and potassium) may be responsible for, at least partial, the intolerance of C. album to high concentration of KCl. In conclusion, C. album could tolerate high concentration of NaCl stress, accompanied by high ability to accumulate Na+ in leaves. These results could contribute to the further investigation of this promising species, e.g., amending saline soil.

Background and Aims: Chenopodium album is well-known as a serious weed and is a salt-tolerant species inhabiting semi-arid and light-saline environments in Xinjiang, China. It produces large amounts of heteromorphic (black and brown) seeds. The primary aims of the present study were to compare the germination characteristics of heteromorphic seeds, the diversity of plant growth and seed proliferation pattern of the resulting plants, and the correlation between NaCl stress and variation of seed heteromorphism., Methods: The phenotypic characters of heteromorphic seeds, e.g. seed morphology, seed mass and total seed protein were determined. The effects of dry storage at room temperature on dormancy behaviour, the germination response of seeds to salinity stress, and the effect of salinity on growth and seed proliferation with plants derived from different seed types were investigated., Key Results: Black and brown seeds differed in seed morphology, mass, total seed protein, dormancy behaviour and salinity tolerance. Brown seeds were large, non-dormant and more salt tolerant, and could germinate rapidly to a high percentage in a wider range of environments; black seeds were salt-sensitive, and a large proportion of seeds were dormant. These characteristics varied between two populations. There was little difference in growth characteristics and seed output of plants produced from the two seed morphs except when plants were subjected to high salinity stress. Plants that suffered higher salinity stress produced more brown (salt-tolerant) seeds., Conclusions: The two seed morphs of C. album exhibited distinct diversity in germination characteristics. There was a significant difference in plant development and seed proliferation pattern from the two types of seeds only when the parent plants were treated with high salinity. In addition, seed heteromorphism of C. album varied between the two populations, and such variation may be attributed, at least in part, to the salinity.

The carry-over effect of sub-lethal herbicides was investigated on the germination of seeds collected from surviving Chenopodium album plants, which had received 1/8, 1/8 twice, 1/8 three times, 1/4, 1/2, 1/1 doses of either pre-emergence ioxynil or post-emergence bentazone in a previous onion (Allium cepa) crop. Seeds were also collected from surviving C. album plants, which had received 1/4, 1/2, 1/1 of either pre-emergence pendimethalin, propachlor and linuron, or 1/8, 1/8 twice, 1/8 three times, 1/4, 1/2, 1/1 of post-emergence ioxynil or linuron in a previous leek (Allium porrum) crop. Seeds of surviving plants were collected and tested for germination at temperature of 5, 10, 15, 20 and 25 degrees C. The effect of different temperatures on the total number of germinated seeds was significant. Germination was minimum at low temperatures (5 degrees C or 10 degrees C). Herbicides did not show any effect on germination of C. album and resulted in the same final germination percentage as seeds collected from the unsprayed control plots.

Production of heat-shock proteins (Hsps) is a key adaptation to acute heat stress and will be important in determining plant responses to climate change. Further, intraspecifc variation in Hsps, which will influence species-level response to global warming, has rarely been examined in naturally occurring plants. To understand intraspecific variation in plant Hsps and its relevance to global warming, we examined Hsp content and thermotolerance in five naturally occurring populations of Chenopodium album L. from contrasting thermal environments grown at low and high temperatures. As expected, Hsp accumulation varied between populations, but this was related more to habitat variability than to mean temperature. Unexpectedly, Hsp accumulation decreased with increasing variability of habitat temperatures. Hsp accumulation also decreased with increased experimental growth temperatures. Physiological thermotolerance was partitioned into basal and induced components. As with Hsps, induced thermotolerance decreased with increasing temperature variability. Thus, populations native to the more stressful habitats, or grown at higher temperatures, had lower Hsp levels and induced thermotolerance, suggesting a greater reliance on basal mechanisms for thermotolerance. These results suggest that future global climate change will differentially impact ecotypes within species, possibly by selecting for increased basal versus inducible thermotolerance.

Determining effects of elevated CO2 on the tolerance of photosynthesis to acute heat-stress (heat wave) is necessary for predicting plant responses to global warming, as photosynthesis is thermolabile and acute heat-stress and atmospheric CO2 will increase in the future. Few studies have examined this, and past results are variable, which may be due to methodological variation. To address this, we grew two C3 and two C4 species at current or elevated CO2 and three different growth temperatures (GT). We assessed photosynthetic thermotolerance in both unacclimated (basal tolerance) and pre-heat-stressed (preHS = acclimated) plants. In C3 species, basal thermotolerance of net photosynthesis (P(n)) was increased in high CO2, but in C4 species, P(n) thermotlerance was decreased by high CO2 (except Zea mays at low GT); CO2 effects in preHS plants were mostly small or absent, though high CO2 was detrimental in one C3 and one C4 species at warmer GT. Though high CO2 generally decreased stomatal conductance, decreases in P(n) during heat stress were mostly due to non-stomatal effects. Photosystem II (PSII) efficiency was often decreased by high CO2 during heat stress, especially at high GT; CO2 effects on post-PSII electron transport were variable. Thus, high CO2 often affected photosynthetic theromotolerance, and the effects varied with photosynthetic pathway, growth temperature, and acclimation state. Most importantly, in heat-stressed plants at normal or warmer growth temperatures, high CO2 may often decrease, or not benefit as expected, tolerance of photosynthesis to acute heat stress. Therefore, interactive effects of elevated CO2 and warmer growth temperatures on acute heat tolerance may contribute to future changes in plant productivity, distribution, and diversity.

Petiole curvature is important in regulating light interception by the leaf. To dissect the determination processes of leaf angle, we irradiated the lamina or petiole of Chenopodium album L. with either one or two spots of actinic light, after dark adaptation. When the abaxial side of the petiole was irradiated with blue light, the petiole curvature increased, and under continuous irradiation, the curvature continued for up to 6 h. The rate of curvature increased with increasing blue light intensity. The curvature induced by irradiation of the abaxial side with blue light ceased when the adaxial side of the petiole was simultaneously irradiated with either blue or red light. When an inhibitor for photosynthesis, 3-(3,4-dichlorophenyl)-1,1-dimethylurea, was applied to the adaxial side of the petiole, the cessation of curvature caused by blue light was only weakly inhibited, while the cessation caused by red light was markedly inhibited. When the adaxial side of the petiole was irradiated alternately with red and far-red light, the far-red light antagonized the cessation of curvature caused by the red light. These results clearly show that the petiole curvature is controlled by two processes, the induction and the cessation of curvature. At least three photoreceptor systems, blue-light receptor, photosynthesis and phytochrome, are involved in the reactions.

Knowledge of soil seed banks is essential to understand the dynamics of plant populations and communities and would greatly benefit from the integration of existing knowledge on ecological correlations of seed size and shape. The present study aims to establish a feasible and meaningful method to describe size-number distributions of seeds in multi-species situations. For that purpose, size-number distributions of seeds with known length, width and thickness were determined by sequential sieving. The most appropriate combination of sieves and seeds dimensions was established, and the adequacy of the power function and the Weibull model to describe size-number distributions of spherical, non-spherical, and all seeds was investigated. We found that the geometric mean of seed length, width and thickness was the most adequate size estimator, providing shape-independent measures of seeds volume directly related to sieves mesh side, and that both the power function and the Weibull model provide high quality descriptions of size-number distributions of spherical, non-spherical, and all seeds. We also found that, in spite of its slightly lower accuracy, the power function is, at this stage, a more trustworthy model to characterize size-number distributions of seeds in soil banks because in some Weibull equations the estimates of the scale parameter were not acceptable.

Background: A Chenopodium album L. biotype surviving in atrazine-treated Serbian corn fields (VC) was compared against atrazine-susceptible (S) and atrazine-resistant (R) standards., Results: Atrazine (2 kg ha(-1)) killed S and VC shoot biomass 15 days after treatment (DAT), but R was only suppressed by 42% and survived 8 kg ha(-1). Atrazine at 2 kg ha(-1) only inhibited VC height by 60% as against 100 and 0% for S and R respectively. Chlorophyll fluorescence (Fv/Fm) and transpiration were insensitive to atrazine in R, but were inhibited by 90 and 100% in S and by 50 and 60% in VC respectively. Decline of Fv/Fm after 2 kg ha(-1) atrazine was stabilized at 3 DAT for the VC biotype., Conclusion: A toxicity mitigation mechanism could have facilitated VC survival in an atrazine-treated field. Further knowledge on this mechanism is needed to establish if surviving VC plants are indicators of atrazine resistance evolution in these Serbian corn fields. Variables related to foliar function provided better detection of weed mechanisms to survive herbicide action than the usual shoot biomass measurements.

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.

Temperature dependence of photoinhibition and photoprotective mechanisms (10-35 degrees C) was investigated for Chenopodium album leaves grown at 25 degrees C under 500 micro mol quanta m(-2) s(-1). The fraction of active photosystem II (PSII) was determined after photoinhibitory treatment at different temperatures in the presence and absence of lincomycin, an inhibitor of chloroplast-encoded protein synthesis. In the absence of lincomycin, leaves were more tolerant to photoinhibition at high (25-35 degrees C) than at low (11-15 degrees C) temperatures. In the presence of lincomycin, the variation in the tolerance to photoinactivation became relatively small. The rate constant of photoinactivation (k(pi)) was stable at 25-35 degrees C and increased by 50% with temperature decrease from 25 to 11 degrees C. The rate constant of recovery of inactivated PSII (k(rec)) was more sensitive to temperature; it was very low at 11 degrees C and increased by an order of magnitude at 35 degrees C. We conclude that the recovery of photoinactivated PSII plays an essential role in photoprotection at 11-35 degrees C. Partitioning of light energy to various photoprotective mechanisms was further analyzed to reveal the factor responsible for k(pi). The fraction of energy utilized in photochemistry was lower at lower temperatures. Although the fraction of heat dissipation increased with decreasing temperatures, the excess energy that is neither utilized by photochemistry nor dissipated by heat dissipation was found to be greater at lower temperatures. The k(pi) value was strongly correlated with the excess energy, suggesting that the excess energy determines the rate of photoinactivation.

Photoinactivation of PSII is thought to be caused by the excessive light energy that is neither used for photosynthetic electron transport nor dissipated as heat. However, the relationship between the photoinactivation rate and excess energy has not been quantitatively evaluated. Chenopodium album L. plants grown under high-light and high-nitrogen (HL-HN) conditions show higher tolerance to photoinactivation and have higher photosynthetic capacity than the high-light and low-nitrogen (HL-LN)- and low-light and high-nitrogen (LL-HN)-grown plants. The rate of photoinactivation in the LL-HN plants was faster than that in the HL-LN, which was similar to that in the HL-HN plants, while the LL-HN and HL-LN plants had similar photosynthetic capacities [Kato et al. (2002b) Funct. Plant Biol. 29: 787]. We quantified partitioning of light energy between the electron transport and heat dissipation at the light intensities ranging from 300 to 1,800 micromol m(-2) s(-1). The maximum electron transport rate was highest in the HL-HN plants, heat dissipation was greatest in the HL-LN plants, and the excess energy, which was neither consumed for electron transport nor dissipated as heat, was greatest in the LL-HN plants. The first-order rate constant of the PSII photoinactivation was proportional to the magnitude of excess energy, with a single proportional constant for all the plants, irrespective of their growth conditions. Thus the excess energy primarily determines the rate of PSII photoinactivation. A large photosynthetic capacity in the HL-HN plants and a large heat dissipation capacity in the HL-LN plants both contribute to the protection of PSII against photoinactivation.

The effect of 2-benzylamino-1,3,5-triazines on photosynthetic electron transport (PET) was measured with thylakoids isolated from atrazine-resistant, wild-type Chenopodium album, and spinach to find novel 1,3,5-triazine herbicides bearing a strong PET inhibition. The PET inhibition assay with Chenopodium (wild-type and resistant), yielded a resistance ratio (R/W = I50 (resistant)/I50 (wild-type)) of 324 for atrazine while for benzylamino-1,3,5-triazine derivatives of diamino-1,3,5-triazines a R/W of 11 to 160 was found. The compounds having a benzylamino group at one of the amino groups in the diamino-1,3,5-triazines have a resistant ratio down to one half to 1/30 of the atrazine value. The average resistance ratio of 21 benzylamino derivatives of monoamino-1,3,5-triazines was found to be about 4.0. The inhibition of 21 benzylamino-1,3,5-triazines assayed with atrazine-resistant Chenopodium thylakoids, indicated by pI50 (R)-values, correlated well with the PET inhibition pI50 (W) of wild-type thylakoids from Chenopodium.