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1.  Selection for low dormancy in annual ryegrass (Lolium rigidum) seeds results in high constitutive expression of a glucose-responsive α-amylase isoform 
Annals of Botany  2012;110(8):1641-1650.
Background and Aims
α-Amylase in grass caryopses (seeds) is usually expressed upon commencement of germination and is rarely seen in dry, mature seeds. A heat-stable α-amylase activity was unexpectedly selected for expression in dry annual ryegrass (Lolium rigidum) seeds during targeted selection for low primary dormancy. The aim of this study was to characterize this constitutive activity biochemically and determine if its presence conferred insensitivity to the germination inhibitors abscisic acid and benzoxazolinone.
Methods
α-Amylase activity in developing, mature and germinating seeds from the selected (low-dormancy) and a field-collected (dormant) population was characterized by native activity PAGE. The response of seed germination and α-amylase activity to abscisic acid and benzoxazolinone was assessed. Using an alginate affinity matrix, α-amylase was purified from dry and germinating seeds for analysis of its enzymatic properties.
Key Results
The constitutive α-amylase activity appeared late during seed development and was mainly localized in the aleurone; in germinating seeds, this activity was responsive to both glucose and gibberellin. It migrated differently on native PAGE compared with the major activities in germinating seeds of the dormant population, but the enzymatic properties of α-amylase purified from the low-dormancy and dormant seeds were largely indistinguishable. Seed imbibition on benzoxazolinone had little effect on the low-dormancy seeds but greatly inhibited germination and α-amylase activity in the dormant population.
Conclusions
The constitutive α-amylase activity in annual ryegrass seeds selected for low dormancy is electrophoretically different from that in germinating seeds and its presence confers insensitivity to benzoxazolinone. The concurrent selection of low dormancy and constitutive α-amylase activity may help to enhance seedling establishment under competitive conditions.
doi:10.1093/aob/mcs213
PMCID: PMC3503496  PMID: 23002268
α-amylase; annual ryegrass; benzoxazolinone; dormancy; germination; Lolium rigidum; seed; xylanase inhibitor protein
2.  Herbicide-resistant weeds: from research and knowledge to future needs 
Evolutionary Applications  2013;6(8):1218-1221.
Synthetic herbicides have been used globally to control weeds in major field crops. This has imposed a strong selection for any trait that enables plant populations to survive and reproduce in the presence of the herbicide. Herbicide resistance in weeds must be minimized because it is a major limiting factor to food security in global agriculture. This represents a huge challenge that will require great research efforts to develop control strategies as alternatives to the dominant and almost exclusive practice of weed control by herbicides. Weed scientists, plant ecologists and evolutionary biologists should join forces and work towards an improved and more integrated understanding of resistance across all scales. This approach will likely facilitate the design of innovative solutions to the global herbicide resistance challenge.
doi:10.1111/eva.12098
PMCID: PMC3901551  PMID: 24478803
agriculture; global crops; herbicide resistance; population genetics; plant adaptation; weed science
3.  Identification of Genetic Elements Associated with EPSPS Gene Amplification 
PLoS ONE  2013;8(6):e65819.
Weed populations can have high genetic plasticity and rapid responses to environmental selection pressures. For example, 100-fold amplification of the 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) gene evolved in the weed species Amaranthus palmeri to confer resistance to glyphosate, the world’s most important herbicide. However, the gene amplification mechanism is unknown. We sequenced the EPSPS gene and genomic regions flanking EPSPS loci in A. palmeri, and searched for mobile genetic elements or repetitive sequences. The EPSPS gene was 10,229 bp, containing 8 exons and 7 introns. The gene amplification likely proceeded through a DNA-mediated mechanism, as introns exist in the amplified gene copies and the entire amplified sequence is at least 30 kb in length. Our data support the presence of two EPSPS loci in susceptible (S) A. palmeri, and that only one of these was amplified in glyphosate-resistant (R) A. palmeri. The EPSPS gene amplification event likely occurred recently, as no sequence polymorphisms were found within introns of amplified EPSPS copies from R individuals. Sequences with homology to miniature inverted-repeat transposable elements (MITEs) were identified next to EPSPS gene copies only in R individuals. Additionally, a putative Activator (Ac) transposase and a repetitive sequence region were associated with amplified EPSPS genes. The mechanism controlling this DNA-mediated amplification remains unknown. Further investigation is necessary to determine if the gene amplification may have proceeded via DNA transposon-mediated replication, and/or unequal recombination between different genomic regions resulting in replication of the EPSPS gene.
doi:10.1371/journal.pone.0065819
PMCID: PMC3677901  PMID: 23762434
4.  Seeds of Brassicaceae weeds have an inherent or inducible response to the germination stimulant karrikinolide 
Annals of Botany  2011;108(5):933-944.
Background and Aims
Karrikinolide (KAR1) is a smoke-derived chemical that can trigger seeds to germinate. A potential application for KAR1 is for synchronizing the germination of weed seeds, thereby enhancing the efficiency of weed control efforts. Yet not all species germinate readily with KAR1, and it is not known whether seemingly non-responsive species can be induced to respond. Here a major agronomic weed family, the Brassicaceae, is used to test the hypothesis that a stimulatory response to KAR1 may be present in physiologically dormant seeds but may not be expressed under all circumstances.
Methods
Seeds of eight Brassicaceae weed species (Brassica tournefortii, Raphanus raphanistrum, Sisymbrium orientale, S. erysimoides, Rapistrum rugosum, Lepidium africanum, Heliophila pusilla and Carrichtera annua) were tested for their response to 1 µm KAR1 when freshly collected and following simulated and natural dormancy alleviation, which included wet–dry cycling, dry after-ripening, cold and warm stratification and a 2 year seed burial trial.
Key Results
Seven of the eight Brassicaceae species tested were stimulated to germinate with KAR1 when the seeds were fresh, and the remaining species became responsive to KAR1 following wet–dry cycling and dry after-ripening. Light influenced the germination response of seeds to KAR1, with the majority of species germinating better in darkness. Germination with and without KAR1 fluctuated seasonally throughout the seed burial trial.
Conclusions
KAR1 responses are more complex than simply stating whether a species is responsive or non-responsive; light and temperature conditions, dormancy state and seed lot all influence the sensitivity of seeds to KAR1, and a response to KAR1 can be induced. Three response types for generalizing KAR1 responses are proposed, namely inherent, inducible and undetected. Given that responses to KAR1 were either inherent or inducible in all 15 seed lots included in this study, the Brassicaceae may be an ideal target for future application of KAR1 in weed management.
doi:10.1093/aob/mcr198
PMCID: PMC3177676  PMID: 21821831
Brassicaceae; butenolide; germination; karrikinolide; karrikins; KAR1; physiological dormancy; seed dormancy; weeds
5.  Prior hydration of Brassica tournefortii seeds reduces the stimulatory effect of karrikinolide on germination and increases seed sensitivity to abscisic acid 
Annals of Botany  2010;105(6):1063-1070.
Background and Aims
The smoke-derived compound karrikinolide (KAR1) shows significant potential as a trigger for the synchronous germination of seeds in a variety of plant-management contexts, from weed seeds in paddocks, to native seeds when restoring degraded lands. Understanding how KAR1 interacts with seed physiology is a necessary precursor to the development of the compound as an efficient and effective management tool. This study tested the ability of KAR1 to stimulate germination of seeds of the global agronomic weed Brassica tournefortii, at different hydration states, to gain insight into how the timing of KAR1 applications in the field should be managed relative to rain events.
Methods
Seeds of B. tournefortii were brought to five different hydration states [equilibrated at 15 % relative humidity (RH), 47 % RH, 96 % RH, fully imbibed, or re-dried to 15 % RH following maximum imbibition] then exposed to 1 nm or 1 µm KAR1 for one of five durations (3 min, 1 h, 24 h, 14 d or no exposure).
Key Results
Dry seeds with no history of imbibition were the most sensitive to KAR1; sensitivity was lower in seeds that were fully imbibed or fully imbibed then re-dried. In addition, reduced sensitivity to KAR1 was associated with an increased sensitivity to exogenously applied abscisic acid (ABA).
Conclusions
Seed water content and history of imbibition were found to significantly influence whether seeds germinate in response to KAR1. To optimize the germination response of seeds, KAR1 should be applied to dry seeds, when sensitivity to ABA is minimized.
doi:10.1093/aob/mcq061
PMCID: PMC2876004  PMID: 20348089
Karrikinolide; karrikins; butenolide; smoke; germination stimulant; seed water content; abscisic acid; ABA; gibberellin; weed; Brassica tournefortii
6.  Selection for low or high primary dormancy in Lolium rigidum Gaud seeds results in constitutive differences in stress protein expression and peroxidase activity 
Journal of Experimental Botany  2010;62(3):1037-1047.
Seed dormancy in wild Lolium rigidum Gaud (annual ryegrass) populations is highly variable and not well characterized at the biochemical level. To identify some of the determinants of dormancy level in these seeds, the proteomes of subpopulations selected for low and high levels of primary dormancy were compared by two-dimensional polyacrylamide gel electrophoresis of extracts from mature, dry seeds. High-dormancy seeds showed higher expression of small heat shock proteins, enolase, and glyoxalase I than the low-dormancy seeds. The functional relevance of these differences in protein expression was confirmed by the fact that high-dormancy seeds were more tolerant to high temperatures imposed at imbibition and had consistently higher glyoxalase I activity over 0–42 d dark stratification. Higher expression of a putative glutathione peroxidase in low-dormancy seeds was not accompanied by higher activity, but these seeds had a slightly more oxidized glutathione pool and higher total peroxidase activity. Overall, these biochemical and physiological differences suggest that L. rigidum seeds selected for low dormancy are more prepared for rapid germination via peroxidase-mediated cell wall weakening, whilst seeds selected for high dormancy are constitutively prepared to survive environmental stresses, even in the absence of stress during seed development.
doi:10.1093/jxb/erq334
PMCID: PMC3022398  PMID: 20974739
Dormancy; glutathione; glyoxalase I; heat shock protein; Lolium rigidum; peroxidase; proteomics; seed
7.  AHAS herbicide resistance endowing mutations: effect on AHAS functionality and plant growth 
Journal of Experimental Botany  2010;61(14):3925-3934.
Twenty-two amino acid substitutions at seven conserved amino acid residues in the acetohydroxyacid synthase (AHAS) gene have been identified to date that confer target-site resistance to AHAS-inhibiting herbicides in biotypes of field-evolved resistant weed species. However, the effect of resistance mutations on AHAS functionality and plant growth has been investigated for only a very few mutations. This research investigates the effect of various AHAS resistance mutations in Lolium rigidum on AHAS functionality and plant growth. The enzyme kinetics of AHAS from five purified L. rigidum populations, each homozygous for the resistance mutations Pro-197-Ala, Pro-197-Arg, Pro-197-Gln, Pro-197-Ser or Trp-574-Leu, were characterized and the pleiotropic effect of three mutations on plant growth was assessed via relative growth rate analysis. All these resistance mutations endowed a herbicide-resistant AHAS and most resulted in higher extractable AHAS activity, with no-to-minor changes in AHAS kinetics. The Pro-197-Arg mutation slightly (but significantly) increased the Km for pyruvate and remarkably increased sensitivity to feedback inhibition by branched chain amino acids. Whereas the Pro-197-Ser and Trp-574-Leu mutations exhibited no significant effects on plant growth, the Pro-197-Arg mutation resulted in lower growth rates. It is clear that, at least in L. rigidum, these five AHAS resistance mutations have no major impact on AHAS functionality and hence probably no plant resistance costs. These results, in part, explain why so many Pro-197 AHAS resistance mutations in AHAS have evolved and why the Pro-197-Ser and the Trp-574-Leu AHAS resistance mutations are frequently found in many weed species.
doi:10.1093/jxb/erq205
PMCID: PMC2935867  PMID: 20627897
AHAS resistance mutation; enzyme kinetics; herbicide resistance; Lolium rigidum; relative growth rate
8.  ABA inhibits germination but not dormancy release in mature imbibed seeds of Lolium rigidum Gaud. 
Journal of Experimental Botany  2009;60(12):3387-3396.
Dormancy release in imbibed annual ryegrass (Lolium rigidum Gaud.) seeds is promoted in the dark but inhibited in the light. The role of abscisic acid (ABA) in inhibition of dormancy release was found to be negligible, compared with its subsequent effect on germination of dormant and non-dormant seeds. Inhibitors of ABA metabolism had the expected effects on seed germination but did not influence ABA concentration, suggesting that they act upon other (unknown) factors regulating dormancy. Although gibberellin (GA) synthesis was required for germination, the influence of exogenous GA on both germination and dormancy release was minor or non-existent. Embryo ABA concentration was the same following treatments to promote (dark stratification) and inhibit (light stratification) dormancy release; exogenous ABA had no effect on this process. However, the sensitivity of dark-stratified seeds to ABA supplied during germination was lower than that of light-stratified seeds. Therefore, although ABA definitely plays a role in the germination of annual ryegrass seeds, it is not the major factor mediating inhibition of dormancy release in imbibed seeds.
doi:10.1093/jxb/erp175
PMCID: PMC2724689  PMID: 19487389
Abscisic acid; dormancy release; fluridone; germination; gibberellins; Lolium rigidum; seed

Results 1-8 (8)