Publications by authors named "Christopher D Todd"

35 Publications

Variation in the post-smolt growth pattern of wild one sea-winter salmon (Salmo salar L.), and its linkage to surface warming in the eastern North Atlantic Ocean.

J Fish Biol 2021 Jan 21;98(1):6-16. Epub 2020 Oct 21.

Department of Health Management, Atlantic Veterinary College, University of Prince Edward Island, Charlottetown, Prince Edward Island, Canada.

Variation in circulus spacing on the scales of wild Atlantic salmon is indicative of changes in body length growth rate. We analyzed scale circulus spacing during the post-smolt growth period for adult one sea-winter salmon (n = 1947) returning to Scotland over the period 1993-2011. The growth pattern of the scales was subjectively and visually categorized according to the occurrence and zonal sequence of three intercirculus spacing criteria ("Slow", "Fast" and "Check" zones). We applied hierarchical time-series cluster analysis to the empirical circulus spacing data, followed by post hoc analysis of significant changes in growth patterns within the 20 identified clusters. Temporal changes in growth pattern frequencies showed significant correlation with sea surface temperature anomalies during the early months of the post-smolt growth season and throughout the Norwegian Sea. Since the turn of the millennium, we observed (a) a marked decrease in the occurrence of continuous Fast growth; (b) increased frequencies of fish showing an extended period of initially Slow growth; and (c) the occurrence of obvious growth Checks or hiatuses. These changes in post-smolt growth pattern were manifest also in decreases in the mean body length attained by the ocean midwinter, as sea surface temperatures have risen.
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http://dx.doi.org/10.1111/jfb.14552DOI Listing
January 2021

Endogenous retroviruses are a source of enhancers with oncogenic potential in acute myeloid leukaemia.

Nat Commun 2020 07 14;11(1):3506. Epub 2020 Jul 14.

Blizard Institute, Barts and The London School of Medicine and Dentistry, QMUL, London, E1 2AT, UK.

Acute myeloid leukemia (AML) is characterised by a series of genetic and epigenetic alterations that result in deregulation of transcriptional networks. One understudied source of transcriptional regulators are transposable elements (TEs), whose aberrant usage could contribute to oncogenic transcriptional circuits. However, the regulatory influence of TEs and their links to AML pathogenesis remain unexplored. Here we identify six endogenous retrovirus (ERV) families with AML-associated enhancer chromatin signatures that are enriched in binding of key regulators of hematopoiesis and AML pathogenesis. Using both locus-specific genetic editing and simultaneous epigenetic silencing of multiple ERVs, we demonstrate that ERV deregulation directly alters the expression of adjacent genes in AML. Strikingly, deletion or epigenetic silencing of an ERV-derived enhancer suppresses cell growth by inducing apoptosis in leukemia cell lines. This work reveals that ERVs are a previously unappreciated source of AML enhancers that may be exploited by cancer cells to help drive tumour heterogeneity and evolution.
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http://dx.doi.org/10.1038/s41467-020-17206-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7360734PMC
July 2020

Transcriptome Analysis Identifies Plasmodiophora brassicae Secondary Infection Effector Candidates.

J Eukaryot Microbiol 2020 05 11;67(3):337-351. Epub 2020 Feb 11.

Department of Biology, University of Saskatchewan, Saskatoon, SK, S7N 5E2, Canada.

Plasmodiophora brassicae (Wor.) is an obligate intracellular plant pathogen affecting Brassicas worldwide. Identification of effector proteins is key to understanding the interaction between P. brassicae and its susceptible host plants. To date, there is very little information available on putative effector proteins secreted by P. brassicae during a secondary infection of susceptible host plants, resulting in root gall production. A bioinformatics pipeline approach to RNA-Seq data from Arabidopsis thaliana (L.) Heynh. root tissues at 17, 20, and 24 d postinoculation (dpi) identified 32 small secreted P. brassicae proteins (SSPbPs) that were highly expressed over this secondary infection time frame. Functional signal peptides were confirmed for 31 of the SSPbPs, supporting the accuracy of the pipeline designed to identify secreted proteins. Expression profiles at 0, 2, 5, 7, 14, 21, and 28 dpi verified the involvement of some of the SSPbPs in secondary infection. For seven of the SSPbPs, a functional domain was identified using Blast2GO and 3D structure analysis and domain functionality was confirmed for SSPbP22, a kinase localized to the cytoplasm and nucleus.
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http://dx.doi.org/10.1111/jeu.12784DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7317818PMC
May 2020

Functional evaluation of transposable elements as enhancers in mouse embryonic and trophoblast stem cells.

Elife 2019 04 23;8. Epub 2019 Apr 23.

Centre for Genomic Health, Life Sciences Institute, Queen Mary University of London, London, United Kingdom.

Transposable elements (TEs) are thought to have helped establish gene regulatory networks. Both the embryonic and extraembryonic lineages of the early mouse embryo have seemingly co-opted TEs as enhancers, but there is little evidence that they play significant roles in gene regulation. Here we tested a set of long terminal repeat TE families for roles as enhancers in mouse embryonic and trophoblast stem cells. Epigenomic and transcriptomic data suggested that a large number of TEs helped to establish tissue-specific gene expression programmes. Genetic editing of individual TEs confirmed a subset of these regulatory relationships. However, a wider survey via CRISPR interference of RLTR13D6 elements in embryonic stem cells revealed that only a minority play significant roles in gene regulation. Our results suggest that a subset of TEs are important for gene regulation in early mouse development, and highlight the importance of functional experiments when evaluating gene regulatory roles of TEs.
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http://dx.doi.org/10.7554/eLife.44344DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6544436PMC
April 2019

There is a direct link between allantoin concentration and cadmium tolerance in Arabidopsis.

Plant Physiol Biochem 2019 Feb 15;135:441-449. Epub 2018 Nov 15.

Department of Biology, University of Saskatchewan, Saskatoon, S7N 5E2, Canada. Electronic address:

Allantoin, an important intermediate of ureide metabolism, has been the subject of investigation recently due to its dual function in nitrogen recycling and abiotic stress response in plants. Allantoin appears to be the dominant ureide accumulating in response to different abiotic stresses, and mutants containing elevated allantoin concentrations exhibit a stress-tolerant phenotype due to limited reactive oxygen species (ROS) generation. Here we describe the involvement of allantoin in stress response and attempt to explain the regulatory mechanism(s) underlying allantoin function in plants. Growth of wild type Col-0 seedlings in the presence of exogenous allantoin improved root elongation in response to Cd treatment. Allantoin treatment of Col-0 seeds increases superoxide dismutase activity causing an enhanced seed germination and seedling growth following Cd exposure. Additionally, allantoinase-overexpressed (ALNox) lines, with lower levels of allantoin, exhibited more susceptibility to Cd treatment than Col-0 Arabidopsis, implying that there is a positive correlation between allantoin concentration and Cd resistance in plants. Growing ABA-insensitive (abi) mutants on allantoin-containing media and comparison between abi mutants and their wild-type backgrounds demonstrated that the potential regulatory function of allantoin does not require ABA at germination but may be ABA-dependent at later stages of seedling growth, suggesting a potential crosstalk between allantoin-mediated stress response and ABA signalling pathway in plants.
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http://dx.doi.org/10.1016/j.plaphy.2018.11.016DOI Listing
February 2019

Precision and accuracy of Dahl-Lea back-calculated smolt lengths from adult scales of Atlantic salmon Salmo salar.

J Fish Biol 2019 Jan 27;94(1):183-186. Epub 2018 Dec 27.

Scottish Oceans Institute, University of St Andrews, St Andrews, UK.

Using tagged and recaptured Atlantic salmon Salmo salar (n = 106) the present analysis shows that the most commonly applied linear back-calculation method for estimating past length, the Dahl-Lea method, resulted in overestimation of the length of large smolts and underestimation of small smolts. A correction equation (y = 0.53x + 6.23) for estimating true smolt length (y) from lengths back-calculated from adult scale measures (x) to account for these systematic discrepancies is proposed.
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http://dx.doi.org/10.1111/jfb.13863DOI Listing
January 2019

Divergent wiring of repressive and active chromatin interactions between mouse embryonic and trophoblast lineages.

Nat Commun 2018 10 10;9(1):4189. Epub 2018 Oct 10.

Blizard Institute, Barts and The London School of Medicine and Dentistry, QMUL, London, E1 2AT, UK.

The establishment of the embryonic and trophoblast lineages is a developmental decision underpinned by dramatic differences in the epigenetic landscape of the two compartments. However, it remains unknown how epigenetic information and transcription factor networks map to the 3D arrangement of the genome, which in turn may mediate transcriptional divergence between the two cell lineages. Here, we perform promoter capture Hi-C experiments in mouse trophoblast (TSC) and embryonic (ESC) stem cells to understand how chromatin conformation relates to cell-specific transcriptional programmes. We find that key TSC genes that are kept repressed in ESCs exhibit interactions between H3K27me3-marked regions in ESCs that depend on Polycomb repressive complex 1. Interactions that are prominent in TSCs are enriched for enhancer-gene contacts involving key TSC transcription factors, as well as TET1, which helps to maintain the expression of TSC-relevant genes. Our work shows that the first developmental cell fate decision results in distinct chromatin conformation patterns establishing lineage-specific contexts involving both repressive and active interactions.
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http://dx.doi.org/10.1038/s41467-018-06666-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6180096PMC
October 2018

Corrigendum to "Allantoin contributes to the stress response in cadmium-treated Arabidopsis roots" [Plant Physiol. Biochem. 119 (October 2017) 103-109].

Plant Physiol Biochem 2018 11 4;132:727. Epub 2018 Oct 4.

Department of Biology, University of Saskatchewan, Saskatoon, S7N 5E2, Canada. Electronic address:

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http://dx.doi.org/10.1016/j.plaphy.2018.09.017DOI Listing
November 2018

Identification of Plasmodiophora brassicae effectors - A challenging goal.

Virulence 2018 ;9(1):1344-1353

a Department of Biology , University of Saskatchewan , Saskatoon , Canada.

Clubroot is an economically important disease affecting Brassica plants worldwide. Plasmodiophora brassicae is the protist pathogen associated with the disease, and its soil-borne obligate parasitic nature has impeded studies related to its biology and the mechanisms involved in its infection of the plant host. The identification of effector proteins is key to understanding how the pathogen manipulates the plant's immune response and the genes involved in resistance. After more than 140 years studying clubroot and P. brassicae, very little is known about the effectors playing key roles in the infection process and subsequent disease progression. Here we analyze the information available for identified effectors and suggest several features of effector genes that can be used in the search for others. Based on the information presented in this review, we propose a comprehensive bioinformatics pipeline for effector identification and provide a list of the bioinformatics tools available for such.
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http://dx.doi.org/10.1080/21505594.2018.1504560DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6177251PMC
April 2019

Transcriptome analysis of response to Plasmodiophora brassicae infection in the Arabidopsis shoot and root.

BMC Genomics 2018 01 5;19(1):23. Epub 2018 Jan 5.

Department of Biology, University of Saskatchewan, Saskatoon, S7N 5E2, Canada.

Background: Clubroot is an important disease caused by the obligate parasite Plasmodiophora brassicae that infects the Brassicaceae. As a soil-borne pathogen, P. brassicae induces the generation of abnormal tissue in the root, resulting in the formation of galls. Root infection negatively affects the uptake of water and nutrients in host plants, severely reducing their growth and productivity. Many studies have emphasized the molecular and physiological effects of the clubroot disease on root tissues. The aim of the present study is to better understand the effect of P. brassicae on the transcriptome of both shoot and root tissues of Arabidopsis thaliana.

Results: Transcriptome profiling using RNA-seq was performed on both shoot and root tissues at 17, 20 and 24 days post inoculation (dpi) of A. thaliana, a model plant host for P. brassicae. The number of differentially expressed genes (DEGs) between infected and uninfected samples was larger in shoot than in root. In both shoot and root, more genes were differentially regulated at 24 dpi than the two earlier time points. Genes that were highly regulated in response to infection in both shoot and root primarily were involved in the metabolism of cell wall compounds, lipids, and shikimate pathway metabolites. Among hormone-related pathways, several jasmonic acid biosynthesis genes were upregulated in both shoot and root tissue. Genes encoding enzymes involved in cell wall modification, biosynthesis of sucrose and starch, and several classes of transcription factors were generally differently regulated in shoot and root.

Conclusions: These results highlight the similarities and differences in the transcriptomic response of above- and below-ground tissues of the model host Arabidopsis following P. brassicae infection. The main transcriptomic changes in root metabolism during clubroot disease progression were identified. An overview of DEGs in the shoot underlined the physiological changes in above-ground tissues following pathogen establishment and disease progression. This study provides insights into host tissue-specific molecular responses to clubroot development and may have applications in the development of clubroot markers for more effective breeding strategies.
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http://dx.doi.org/10.1186/s12864-017-4426-7DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5756429PMC
January 2018

Exogenous allantoin increases Arabidopsis seedlings tolerance to NaCl stress and regulates expression of oxidative stress response genes.

J Plant Physiol 2018 Feb 7;221:43-50. Epub 2017 Dec 7.

University of Saskatchewan, Department of Biology, 112 Science Place, Saskatoon, SK, S7N5E2, Canada. Electronic address:

Allantoin is a nitrogenous compound derived from purine catabolism that contributes to nitrogen recycling in plants. Accumulation of allantoin in plant tissues and a potential role in protection of plants from abiotic stress conditions has been identified. The present work shows that application of exogenous allantoin increased stress tolerance of Arabidopsis seedlings when germinated on, or subjected to the media containing NaCl. Allantoin-induced tolerance to NaCl stress was associated with decreased production of superoxide and hydrogen peroxide in seedlings. To understand the molecular mechanism, the effect of exogenous allantoin treatment on expression of several stress-related genes was investigated. Exogenous allantoin altered the expression of several antioxidant encoding genes and upregulated the expression of two genes involved in oxidative stress tolerance, SOS1 and RCD1, in the presence or absence of NaCl. Allantoin increased the NaCl tolerance of abscisic acid insensitive mutants, suggesting that it can function independently of abscisic acid signaling. These results provide additional evidence for the role of allantoin in enhancing plants tolerance to oxidative stress.
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http://dx.doi.org/10.1016/j.jplph.2017.11.011DOI Listing
February 2018

Allantoin contributes to the stress response in cadmium-treated Arabidopsis roots.

Plant Physiol Biochem 2017 Oct 17;119:103-109. Epub 2017 Aug 17.

Department of Biology, University of Saskatchewan, Saskatoon, S7N 5E2, Canada. Electronic address:

Ureides are nitrogen-rich compounds, derived from purine catabolism. A dual role for ureides, and for allantoin in particular, in both nitrogen recycling and the abiotic stress response has been recently identified. Previous work on the effect of allantoin on cadmium (Cd)-exposed Arabidopsis revealed that high concentration of allantoin in allantoinase-negative mutant (aln-3) leaves alleviates Cd toxicity via inducing antioxidant mechanisms in these plants. In the present study, we evaluate whether allantoin has a similar protective role in roots. Both wild type and aln-3 roots contain higher amounts of internal Cd compared to leaves. Likewise, aln-3 roots are more resistant to Cd, reflected in fresh and dry weight, and stimulated antioxidant enzyme activity, including superoxide dismutase (SOD) and catalase (CAT), resulting in lower reactive oxygen species concentration. In contrast with wild-type leaves, high levels of Cd in Col-0 roots reduces transcript abundance of uricase, leading to a significant decline in allantoin level of treated roots at 1000 and 1500 μM CdCl. This metabolite change is also accompanied by decreasing the activity of antioxidant enzymes (SOD and CAT). Additionally, contrary to wild-type leaves, root genotype has a significant effect on CAT activity under Cd treatment, suggesting the possible different sources of damage and oxidative stress response in these two tissues.
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http://dx.doi.org/10.1016/j.plaphy.2017.08.012DOI Listing
October 2017

TET-dependent regulation of retrotransposable elements in mouse embryonic stem cells.

Genome Biol 2016 11 18;17(1):234. Epub 2016 Nov 18.

Blizard Institute, Barts and The London School of Medicine and Dentistry, QMUL, London, E1 2AT, UK.

Background: Ten-eleven translocation (TET) enzymes oxidise DNA methylation as part of an active demethylation pathway. Despite extensive research into the role of TETs in genome regulation, little is known about their effect on transposable elements (TEs), which make up nearly half of the mouse and human genomes. Epigenetic mechanisms controlling TEs have the potential to affect their mobility and to drive the co-adoption of TEs for the benefit of the host.

Results: We performed a detailed investigation of the role of TET enzymes in the regulation of TEs in mouse embryonic stem cells (ESCs). We find that TET1 and TET2 bind multiple TE classes that harbour a variety of epigenetic signatures indicative of different functional roles. TETs co-bind with pluripotency factors to enhancer-like TEs that interact with highly expressed genes in ESCs whose expression is partly maintained by TET2-mediated DNA demethylation. TETs and 5-hydroxymethylcytosine (5hmC) are also strongly enriched at the 5' UTR of full-length, evolutionarily young LINE-1 elements, a pattern that is conserved in human ESCs. TETs drive LINE-1 demethylation, but surprisingly, LINE-1s are kept repressed through additional TET-dependent activities. We find that the SIN3A co-repressive complex binds to LINE-1s, ensuring their repression in a TET1-dependent manner.

Conclusions: Our data implicate TET enzymes in the evolutionary dynamics of TEs, both in the context of exaptation processes and of retrotransposition control. The dual role of TET action on LINE-1s may reflect the evolutionary battle between TEs and the host.
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http://dx.doi.org/10.1186/s13059-016-1096-8DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5116139PMC
November 2016

Allantoin Increases Cadmium Tolerance in Arabidopsis via Activation of Antioxidant Mechanisms.

Plant Cell Physiol 2016 Dec 13;57(12):2485-2496. Epub 2016 Oct 13.

Department of Biology, University of Saskatchewan, Saskatoon, SK, S7N 5E2, Canada

Plants apply various molecular, physiological and morphological strategies in response to undesirable environmental conditions. One of the possible responses which may contribute to surviving stressful conditions is the accumulation of ureides. Ureides are recognized as important nitrogen-rich compounds involved in recycling nitrogen in plants to support growth and reproduction. Amongst them, allantoin not only serves as a transportable nitrogen-rich compound, but has also been suggested to protect plants from abiotic stresses via minimizing oxidative damage. This work focuses on the effect of cadmium (Cd) on ureide metabolism in Arabidopsis, in order to clarify the potential role of allantoin in plant tolerance to heavy metals. In response to Cd treatment, allantoin levels increase in Arabidopsis thaliana, ecotype Col-0, due to reduced allantoinase (ALN) gene expression and enzyme activity. This coincides with increases in uricase (UO) transcripts. UO and ALN encode the enzymes for the production and degradation of allantoin, respectively. ALN-negative aln-3 Arabidopsis mutants with elevated allantoin levels demonstrate resistance to soil-applied CdCl, up to 1,500 μM. Although aln-3 mutants take up and store more Cd within their leaf tissue, they contain less damaging superoxide radicals. The protective mechanism of aln-3 mutants appears to involve enhancing the activity of antioxidant enzymes such as superoxide dismutase and ascorbate peroxidase.
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http://dx.doi.org/10.1093/pcp/pcw162DOI Listing
December 2016

Ureide metabolism under abiotic stress in Arabidopsis thaliana.

J Plant Physiol 2016 Jul 21;199:87-95. Epub 2016 May 21.

Department of Biology, University of Saskatchewan, Saskatoon, SK, S7N 5E2, Canada. Electronic address:

Ureides are nitrogenous compounds derived from purine catabolism which contribute to nitrogen recycling in plants. Accumulation of ureide compounds has been reported in a number of plant species under stress conditions, suggesting their involvement in plants' response to stress. In this research a biochemical and molecular approach was applied to address the ureide accumulation under abiotic stress conditions in Arabidopsis thaliana. Ureide concentration and changes in expression of ureide metabolic genes were examined in response to drought, NaCl and mannitol treatments. Additionally, an Arabidopsis allantoinase (ALN) mutant with constitutive accumulation of a ureide compound, allantoin, was used to investigate the impact of high levels of this compound on drought and NaCl stress responses. In the leaf tissue of adult plants allantoin accumulated in response to soil drying. Transcription of urate oxidase (UO), involved in allantoin production, was highly up-regulated under the same conditions. Allantoin and allantoate also accumulated in seedlings following treatment with NaCl or mannitol. aln mutants with enhanced levels of allantoin exhibited higher tolerance to drought and NaCl. Hydrogen peroxide and superoxide did not accumulate in the aln mutant leaves to the same degree in response to drought when compared to the wild-type. Our results suggest that ureide metabolism and accumulation contribute to the abiotic stress response which is regulated, at least in part, at the transcriptional level. Higher concentrations of allantoin in the mutant elevates abiotic stress tolerance, possibly by reducing oxidative damage.
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http://dx.doi.org/10.1016/j.jplph.2016.05.011DOI Listing
July 2016

Determination of Photoperiod-Sensitive Phase in Chickpea (Cicer arietinum L.).

Front Plant Sci 2016 11;7:478. Epub 2016 Apr 11.

Crop Development Centre/Department of Plant Sciences, University of Saskatchewan Saskatoon, SK, Canada.

Photoperiod is one of the major environmental factors determining time to flower initiation and first flower appearance in plants. In chickpea, photoperiod sensitivity, expressed as delayed to flower under short days (SD) as compared to long days (LD), may change with the growth stage of the crop. Photoperiod-sensitive and -insensitive phases were identified by experiments in which individual plants were reciprocally transferred in a time series from LD to SD and vice versa in growth chambers. Eight chickpea accessions with differing degrees of photoperiod sensitivity were grown in two separate chambers, one of which was adjusted to LD (16 h light/8 h dark) and the other adjusted to SD (10 h light/14 h dark), with temperatures of 22/16°C (12 h light/12 h dark) in both chambers. The accessions included day-neutral (ICCV 96029 and FLIP 98-142C), intermediate (ICC 15294, ICC 8621, ILC 1687, and ICC 8855), and photoperiod-sensitive (CDC Corinne and CDC Frontier) responses. Control plants were grown continuously under the respective photoperiods. Reciprocal transfers of plants between the SD and LD photoperiod treatments were made at seven time points after sowing, customized for each accession based on previous data. Photoperiod sensitivity was detected in intermediate and photoperiod-sensitive accessions. For the day-neutral accession, ICCV 96029, there was no significant difference in the number of days to flowering of the plants grown under SD and LD as well as subsequent transfers. In photoperiod-sensitive accessions, three different phenological phases were identified: a photoperiod-insensitive pre-inductive phase, a photoperiod-sensitive inductive phase, and a photoperiod-insensitive post-inductive phase. The photoperiod-sensitive phase extends after flower initiation to full flower development. Results from this research will help to develop cultivars with shorter pre-inductive photoperiod-insensitive and photoperiod-sensitive phases to fit to regions with short growing seasons.
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http://dx.doi.org/10.3389/fpls.2016.00478DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4826865PMC
May 2016

Arabidopsis AIR12 influences root development.

Physiol Mol Biol Plants 2015 Oct 30;21(4):479-89. Epub 2015 Sep 30.

Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, SK S7N 5E2 Canada.

Arabidopsis AUXIN INDUCED IN ROOTS (AIR 12) is a predicted to encode a glycosylphosphatidylinositol tail anchored protein. It has been associated with extracellular redox processes, but little is known about its physiological role. An air12 mutant line demonstrated increased germination rates in the presence of a range of abiotic stress factors and hormones, but not in the presence of ABA. Disruption of AIR12 also affected primary and lateral root development and was linked to changes in root catalase activity and superoxide production. We suggest AIR12 is an extracellular constituent linking both hormone and reactive oxygen signaling in plants.
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http://dx.doi.org/10.1007/s12298-015-0323-1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4646869PMC
October 2015

Physiological implications of arginine metabolism in plants.

Front Plant Sci 2015 30;6:534. Epub 2015 Jul 30.

Laboratory of Plant Physiology and Biochemistry, Department of Biology, University of Konstanz , Konstanz, Germany.

Nitrogen is a limiting resource for plant growth in most terrestrial habitats since large amounts of nitrogen are needed to synthesize nucleic acids and proteins. Among the 21 proteinogenic amino acids, arginine has the highest nitrogen to carbon ratio, which makes it especially suitable as a storage form of organic nitrogen. Synthesis in chloroplasts via ornithine is apparently the only operational pathway to provide arginine in plants, and the rate of arginine synthesis is tightly regulated by various feedback mechanisms in accordance with the overall nutritional status. While several steps of arginine biosynthesis still remain poorly characterized in plants, much wider attention has been paid to inter- and intracellular arginine transport as well as arginine-derived metabolites. A role of arginine as alternative source besides glutamate for proline biosynthesis is still discussed controversially and may be prevented by differential subcellular localization of enzymes. Apparently, arginine is a precursor for nitric oxide (NO), although the molecular mechanism of NO production from arginine remains unclear in higher plants. In contrast, conversion of arginine to polyamines is well documented, and in several plant species also ornithine can serve as a precursor for polyamines. Both NO and polyamines play crucial roles in regulating developmental processes as well as responses to biotic and abiotic stress. It is thus conceivable that arginine catabolism serves on the one hand to mobilize nitrogen storages, while on the other hand it may be used to fine-tune development and defense mechanisms against stress. This review summarizes the recent advances in our knowledge about arginine metabolism, with a special focus on the model plant Arabidopsis thaliana, and pinpoints still unresolved critical questions.
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http://dx.doi.org/10.3389/fpls.2015.00534DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4520006PMC
August 2015

A series of TA-based and zero-background vectors for plant functional genomics.

PLoS One 2013 29;8(3):e59576. Epub 2013 Mar 29.

Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Chinese Academy of Science, Kunming, Yunna, China.

With the sequencing of genomes from many organisms now complete and the development of high-throughput sequencing, life science research has entered the functional post-genome era. Therefore, deciphering the function of genes and how they interact is in greater demand. To study an unknown gene, the basic methods are either overexpression or gene knockout by creating transgenic plants, and gene construction is usually the first step. Although traditional cloning techniques using restriction enzymes or a site-specific recombination system (Gateway or Clontech cloning technology) are highly useful for efficiently transferring DNA fragments into destination plasmids, the process is time consuming and expensive. To facilitate the procedure of gene construction, we designed a TA-based cloning system in which only one step was needed to subclone a DNA fragment into vectors. Such a cloning system was developed from the pGreen binary vector, which has a minimal size and facilitates construction manipulation, combined with the negative selection marker gene ccdB, which has the advantages of eliminating the self-ligation background and directly enabling high-efficiency TA cloning technology. We previously developed a set of transient and stable transformation vectors for constitutive gene expression, gene silencing, protein tagging, subcellular localization analysis and promoter activity detection. Our results show that such a system is highly efficient and serves as a high-throughput platform for transient or stable transformation in plants for functional genome research.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0059576PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3612078PMC
September 2013

Comparative proteome analyses reveal that nitric oxide is an important signal molecule in the response of rice to aluminum toxicity.

J Proteome Res 2013 Mar 6;12(3):1316-30. Epub 2013 Feb 6.

School of Life Sciences, Jiangsu Key Laboratory for Eco-Agriculture Biotechnology around Hongze Lake, Huaiyin Normal University, Huai'an223300, China.

Acidic soils inhibit crop yield and reduce grain quality. One of the major contributing factors to acidic soil is the presence of soluble aluminum (Al(3+)) ions, but the mechanisms underlying plant responses to Al(3+) toxicity remain elusive. Nitric oxide (NO) is an important messenger and participates in various plant physiological responses. Here, we demonstrate that Al(3+) induced an increase of NO in rice seedlings; adding exogenous NO alleviated the Al(3+) toxicity related to rice growth and photosynthetic capacity, effects that could be reversed by suppressing NO metabolism. Comparative proteomic analyses successfully identified 92 proteins that showed differential expression after Al(3+) or NO treatment. In particular, some of the proteins are involved in reactive oxygen species (ROS) and reactive nitrogen species (RNS) metabolism. Further analyses confirmed that NO treatment reduced Al(3+)-induced ROS and RNS toxicities by increasing the activities and protein expression of antioxidant enzymes, as well as S-nitrosoglutathione reductase (GSNOR). Suppressing GSNOR enzymatic activity aggravated Al(3+) damage to rice and increased the accumulation of RNS. NO treatment altered the expression of proteins associated with cell wall synthesis, cell division and cell structure, calcium signaling and defense responses. On the basis of these results, we propose that NO activates multiple pathways that enhance rice adaptation to Al(3+) toxicity. Such findings may be applicable to crop engineering to enhance yield and improve stress tolerance.
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http://dx.doi.org/10.1021/pr300971nDOI Listing
March 2013

Impact of parasites on salmon recruitment in the Northeast Atlantic Ocean.

Proc Biol Sci 2013 Jan 7;280(1750):20122359. Epub 2012 Nov 7.

Department of Zoology, University of Otago, Dunedin, New Zealand.

Parasites may have large effects on host population dynamics, marine fisheries and conservation, but a clear elucidation of their impact is limited by a lack of ecosystem-scale experimental data. We conducted a meta-analysis of replicated manipulative field experiments concerning the influence of parasitism by crustaceans on the marine survival of Atlantic salmon (Salmo salar L.). The data include 24 trials in which tagged smolts (totalling 283 347 fish; 1996-2008) were released as paired control and parasiticide-treated groups into 10 areas of Ireland and Norway. All experimental fish were infection-free when released into freshwater, and a proportion of each group was recovered as adult recruits returning to coastal waters 1 or more years later. Treatment had a significant positive effect on survival to recruitment, with an overall effect size (odds ratio) of 1.29 that corresponds to an estimated loss of 39 per cent (95% CI: 18-55%) of adult salmon recruitment. The parasitic crustaceans were probably acquired during early marine migration in areas that host large aquaculture populations of domesticated salmon, which elevate local abundances of ectoparasitic copepods-particularly Lepeophtheirus salmonis. These results provide experimental evidence from a large marine ecosystem that parasites can have large impacts on fish recruitment, fisheries and conservation.
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http://dx.doi.org/10.1098/rspb.2012.2359DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3574446PMC
January 2013

Carbon monoxide enhances the chilling tolerance of recalcitrant Baccaurea ramiflora seeds via nitric oxide-mediated glutathione homeostasis.

Free Radic Biol Med 2012 Aug 7;53(4):710-20. Epub 2012 Jun 7.

Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan 650201, China.

Both carbon monoxide (CO) and nitric oxide (NO) play fundamental roles in plant responses to environmental stress. Glutathione (GSH) homeostasis through the glutathione-ascorbate cycle regulates the cellular redox status and protects the plant from damage due to reactive oxygen species (ROS) or reactive nitrogen species (RNS). Most recalcitrant seeds are sensitive to chilling stress, but the roles of and cross talk among CO, NO, ROS, and GSH in recalcitrant seeds under low temperature are not well understood. Here, we report that the germination of recalcitrant Baccaurea ramiflora seeds shows sensitivity to chilling stress, but application of exogenous CO or NO markedly increased GSH accumulation, enhanced the activities of antioxidant enzymes involved in the glutathione-ascorbate cycle, decreased the content of H(2)O(2) and RNS, and improved the tolerance of seeds to low-temperature stress. Compared to orthodox seeds such as maize, only transient accumulation of CO and NO was induced and only a moderate increase in GSH was shown in the recalcitrant B. ramiflora seeds. Exogenous CO or NO treatment further increased the GSH accumulation and S-nitrosoglutathione reductase (GSNOR) activity in B. ramiflora seeds under chilling stress. In contrast, suppressing CO or NO generation, removing GSH, or blocking GSNOR activity resulted in increases in ROS and RNS and impaired the germination of CO- or NO-induced seeds under chilling stress. Based on these results, we propose that CO acts as a novel regulator to improve the tolerance of recalcitrant seeds to low temperatures through NO-mediated glutathione homeostasis.
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http://dx.doi.org/10.1016/j.freeradbiomed.2012.05.042DOI Listing
August 2012

Comparative proteomic analysis of the thermotolerant plant Portulaca oleracea acclimation to combined high temperature and humidity stress.

J Proteome Res 2012 Jul 7;11(7):3605-23. Epub 2012 Jun 7.

Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Institute of Tibet Plateau Research, Chinese Academy of Science, Kunming 650204, China.

Elevated temperature and humidity are major environmental factors limiting crop yield and distribution. An understanding of the mechanisms underlying plant tolerance to high temperature and humidity may facilitate the development of cultivars adaptable to warm or humid regions. Under conditions of 90% humidity and 35 °C, the thermotolerant plant Portulaca oleracea exhibits excellent photosynthetic capability and relatively little oxidative damage. To determine the proteomic response that occurs in leaves of P. oleracea following exposure to high temperature and high humidity, a proteomic approach was performed to identify protein changes. A total of 51 differentially expressed proteins were detected and characterized functionally and structurally; these identified proteins were involved in various functional categories, mainly including material and energy metabolism, the antioxidant defense responses, protein destination and storage, and transcriptional regulation. The subset of antioxidant defense-related proteins demonstrated marked increases in activity with exposure to heat and humidity, which led to lower accumulations of H(2)O(2) and O(2)(-) in P. oleracea compared with the thermosensitive plant Arabidopsis thaliana. The quickly accumulations of proline content and heat-shock proteins, and depleting abscisic acid (ABA) via increasing ABA-8'-hydroxylase were also found in P. oleracea under stress conditions, that resulted into greater stomata conductance and respiration rates. On the basis of these findings, we propose that P. oleracea employs multiple strategies to enhance its adaptation to high-temperature and high-humidity conditions.
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http://dx.doi.org/10.1021/pr300027aDOI Listing
July 2012

Four allantoinase genes are expressed in nitrogen-fixing soybean.

Plant Physiol Biochem 2012 May 10;54:149-55. Epub 2012 Mar 10.

Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, SK S7N 5E2, Canada.

Soybean (Glycine max L. [Merr]) plants export nitrogen from the nodules as ureides during symbiotic biological nitrogen fixation. Ureides also play a role as nitrogen storage compounds in the seeds and are broken down in germinating seedlings. In this work we identified four soybean genes encoding allantoinase (E.C. 3.5.2.5), an enzyme involved in both ureide production in nodules and ureide catabolism in leaves and other sink tissues. We examined ureide content, allantoinase enzyme activity and expression of these genes, which we term GmALN1 through GmALN4, in germinating seedlings and in vegetative tissues from 45 day old soybean plants. GmALN1 and GmALN2 transcripts were measured in all tissues, but similar levels of expression of GmALN3 and GmALN4 was only observed in nodules. The soybean allantoinase genes seem to have arisen through tandem gene duplication followed by a whole genome duplication. We looked for evidence of the tandem duplication in common bean (Phaseolus vulgaris L.) and present evidence that it occured sometime in the bean lineage before these two species diverged, but before soybean became a tetraploid.
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http://dx.doi.org/10.1016/j.plaphy.2012.03.002DOI Listing
May 2012

N-3-oxo-decanoyl-L-homoserine-lactone activates auxin-induced adventitious root formation via hydrogen peroxide- and nitric oxide-dependent cyclic GMP signaling in mung bean.

Plant Physiol 2012 Feb 2;158(2):725-36. Epub 2011 Dec 2.

Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Institute of Tibet Plateau Research at Kunming, Chinese Academy of Sciences, Kunming 650204, China.

N-Acyl-homoserine-lactones (AHLs) are bacterial quorum-sensing signaling molecules that regulate population density. Recent evidence demonstrates their roles in plant defense responses and root development. Hydrogen peroxide (H(2)O(2)), nitric oxide (NO), and cyclic GMP (cGMP) are essential messengers that participate in various plant physiological processes, but how these messengers modulate the plant response to N-acyl-homoserine-lactone signals remains poorly understood. Here, we show that the N-3-oxo-decanoyl-homoserine-lactone (3-O-C10-HL), in contrast to its analog with an unsubstituted branch chain at the C3 position, efficiently stimulated the formation of adventitious roots and the expression of auxin-response genes in explants of mung bean (Vigna radiata) seedlings. This response was mimicked by the exogenous application of auxin, H(2)O(2), NO, or cGMP homologs but suppressed by treatment with scavengers or inhibitors of H(2)O(2), NO, or cGMP metabolism. The 3-O-C10-HL treatment enhanced auxin basipetal transport; this effect could be reversed by treatment with H(2)O(2) or NO scavengers but not by inhibitors of cGMP synthesis. Inhibiting 3-O-C10-HL-induced H(2)O(2) or NO accumulation impaired auxin- or 3-O-C10-HL-induced cGMP synthesis; however, blocking cGMP synthesis did not affect auxin- or 3-O-C10-HL-induced H(2)O(2) or NO generation. Additionally, cGMP partially rescued the inhibitory effect of H(2)O(2) or NO scavengers on 3-O-C10-HL-induced adventitious root development and auxin-response gene expression. These results suggest that 3-O-C10-HL, unlike its analog with an unmodified branch chain at the C3 position, can accelerate auxin-dependent adventitious root formation, possibly via H(2)O(2)- and NO-dependent cGMP signaling in mung bean seedlings.
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http://dx.doi.org/10.1104/pp.111.185769DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3271762PMC
February 2012

The mitochondrial connection: Arginine degradation versus arginine conversion to nitric oxide.

Plant Signal Behav 2008 Dec;3(12):1106-8

Biochemistry Department; University of Missouri; Columbia, Missouri USA.

Arg catabolism to cytoplasmic urea and glutamate is initiated by two mitochondrial enzymes, arginase and ornithine aminotransferase. Mutation of either enzyme leads to Arg sensitivity, and at least in the former, an arginine-induced seedling morphology similar to exogenous auxin treatment. We reported that single mutants lacking either of two arginase isozymes exhibited more NO accumulation and efflux, and increased responses to auxin (measured by DR5 reporter expression and auxin-induced lateral roots). We discuss evidence for stimulation of NO by arginine, either directly, or via polyamines derived from arginine. We favor the "direct" route because mitochondria are sites of NO 'hot spots,' and the location of arginine-degrading enzymes and the NO-associated protein1. The polyamine "branch" invokes more than one cell compartment, at least two intermediates (polyamines and H(2)O(2)) between Arg and NO, and is not consistent with enhanced lateral root formation in arginine decarboxylase mutants. Genetic tools are at our disposal to test the two possible routes of arginine-derived NO.
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http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2634469PMC
http://dx.doi.org/10.4161/psb.3.12.7007DOI Listing
December 2008

Arginase-negative mutants of Arabidopsis exhibit increased nitric oxide signaling in root development.

Plant Physiol 2008 Aug 20;147(4):1936-46. Epub 2008 Jun 20.

Biochemistry Department, University of Missouri, Columbia, Missouri 65211, USA.

Mutation of either arginase structural gene (ARGAH1 or ARGAH2 encoding arginine [Arg] amidohydrolase-1 and -2, respectively) resulted in increased formation of lateral and adventitious roots in Arabidopsis (Arabidopsis thaliana) seedlings and increased nitric oxide (NO) accumulation and efflux, detected by the fluorogenic traps 3-amino,4-aminomethyl-2',7'-difluorofluorescein diacetate and diamino-rhodamine-4M, respectively. Upon seedling exposure to the synthetic auxin naphthaleneacetic acid, NO accumulation was differentially enhanced in argah1-1 and argah2-1 compared with the wild type. In all genotypes, much 3-amino,4-aminomethyl-2',7'-difluorofluorescein diacetate fluorescence originated from mitochondria. The arginases are both localized to the mitochondrial matrix and closely related. However, their expression levels and patterns differ: ARGAH1 encoded the minor activity, and ARGAH1-driven beta-glucuronidase (GUS) was expressed throughout the seedling; the ARGAH2::GUS expression pattern was more localized. Naphthaleneacetic acid increased seedling lateral root numbers (total lateral roots per primary root) in the mutants to twice the number in the wild type, consistent with increased internal NO leading to enhanced auxin signaling in roots. In agreement, argah1-1 and argah2-1 showed increased expression of the auxin-responsive reporter DR5::GUS in root tips, emerging lateral roots, and hypocotyls. We propose that Arg, or an Arg derivative, is a potential NO source and that reduced arginase activity in the mutants results in greater conversion of Arg to NO, thereby potentiating auxin action in roots. This model is supported by supplemental Arg induction of adventitious roots and increased NO accumulation in argah1-1 and argah2-1 versus the wild type.
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http://dx.doi.org/10.1104/pp.108.121459DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2492630PMC
August 2008

Analysis of Arabidopsis arginase gene transcription patterns indicates specific biological functions for recently diverged paralogs.

Plant Mol Biol 2008 Jul 19;67(4):429-40. Epub 2008 Apr 19.

Department of Biological Sciences, University of Alberta, CW 405 BioSci, Univ Alberta, Edmonton, AB, Canada.

The detailed expression patterns of transcripts of two Arabidopsis arginase genes, ARGAH1 and ARGAH2, have not been previously described, and phylogenetic analysis suggests that they diverged independently of duplication events in other lineages. Therefore, we used beta-glucuronidase reporter fusions and quantitative reverse-transcriptase PCR to analyze tissue-specific expression of ARGAH1 and ARGAH2 during Arabidopsis development, and in response to the availability of nutrients and exposure to methyl jasmonate (MeJA). We demonstrated tissue-specific transcript expression and enzyme activity in pollen for ARGAH1, but not ARGAH2. Conversely, we demonstrated MeJA-inducibility of ARGAH2, but not ARGAH1. In addition, we used microarrays to identify genes for which transcript abundance following MeJA treatment differed in wild type and ARGAH2 mutants. These ARGAH2 and MeJA responsive genes included a putative pathogenesis-related protein pathogenesis response-1 (At2g14610), and a gene of unknown function (At5g03090). Interestingly, these genes had opposite responses to the loss of ARGAH2, suggesting multiple downstream effects of arginase activity, following MeJA treatment. These results, and the variety and complexity of expression patterns of ARGAH1 and ARGAH2 transcript expression and their related reporter gene fusions that we observed point to multiple functions of arginase genes in Arabidopsis, some of which have resulted through a sub-functionalization not shared by all angiosperms.
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http://dx.doi.org/10.1007/s11103-008-9336-2DOI Listing
July 2008

Arabidopsis mitochondria have two basic amino acid transporters with partially overlapping specificities and differential expression in seedling development.

Biochim Biophys Acta 2006 Sep-Oct;1757(9-10):1277-83. Epub 2006 Apr 7.

Department of Pharmaco-Biology, Laboratory of Biochemistry and Molecular Biology, University of Bari and CNR Institute of Biomembranes and Bioenergetics, Via E. Orabona 4, 70125 Bari, Italy.

To shed light on the metabolic role of two mitochondrial transporters for basic amino acids in Arabidopsis, we compared their functional properties in liposomes and expression during germination. Recombinant and purified BAC2, as previously reported for BAC1, transported various basic L-amino acids upon reconstitution in phospholipid vesicles. Both displayed highest affinity for arginine with similar Km and Vmax. However, BAC2 transported citrulline for which BAC1 had little or no affinity. Furthermore, BAC2 was less stereospecific than BAC1, transporting D-arginine and D-lysine at significant rates, and displayed a striking alkaline pH optimum (pH 8.0) whereas BAC1 activity was unaltered from pH 7.0 to 9.0. By semi-quantitative RT-PCR BAC1 transcript levels were found to be higher than those of BAC2 in germinated seeds. However, BAC2 expression transiently increased 2 days after germination. Disruption of the Arabidopsis arginase structural genes (ARGAH1 or ARGAH2) accentuated the increases of transcript levels of BAC1 at germination and of BAC2 2 days after germination and from 6 days on. Early expression of BAC1 and BAC2 is consistent with the delivery of arginine, released from seed reserves, to mitochondrial arginase and the export of ornithine. Increase of BAC2 transcript levels later in seedling development is consistent with roles in NO, polyamine or proline metabolism--processes involving arginine, citrulline and/or ornithine.
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http://dx.doi.org/10.1016/j.bbabio.2006.03.025DOI Listing
December 2006

AtAAH encodes a protein with allantoate amidohydrolase activity from Arabidopsis thaliana.

Planta 2006 Apr 22;223(5):1108-13. Epub 2006 Feb 22.

Department of Biochemistry and Interdisciplinary Plant Group, University of Missouri, 117 Schweitzer Hall, Columbia, 65211, USA.

We report the identification and cloning of an allantoate amidohydrolase (allantoate deiminase, EC 3.5.3.9) cDNA from Arabidopsis thaliana (L.) Heynh. This sequence, which we term Arabidopsis thaliana Allantoate Amidohydrolase (AtAAH), was shown to be functional by complementation of Saccharomyces cerevisiae dal2 mutants, blocked in allantoate degradation. Following transfer to a medium containing allantoin as the sole nitrogen source, Ataah T-DNA insertion mutants were severely impaired and eventually died. Ataah mutants demonstrated higher allantoate levels than wild-type plants in the presence and absence of exogenous ureides, supporting a block in allantoate catabolism. AtAAH transcript was detected in all tissues examined by RT-PCR, consistent with a function in purine turnover in Arabidopsis. To our knowledge this is the first allantoate amidohydrolase gene identified in any plant species.
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http://dx.doi.org/10.1007/s00425-006-0236-xDOI Listing
April 2006