Arabidopsis Research Roundup: November 1st.

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Published on: November 1, 2017

This weeks Research Roundup includes three research and two methods papers. Firstly is work from the O’Connor and Leyser groups at SLCU that investigates the diversity of function in PIN auxin transporters between monocots and dicots. Secondly research from the Kover lab at the University of Bath has characterised the photosynthetic contribution of the inflorescence stem whilst the third paper is from the Bill Finch-Savage at the University of Warwick and looks at the effect of temperature on seed dormancy. Finally are two methods paper from the University of Warwick and Leeds that introduce protocols for the imaging of either the endoplasmic reticulum or the ultrastructure of pollen tubes.


O’Connor DL, Elton S, Ticchiarelli F, Hsia MM, Vogel JP, Leyser O (2017) Cross-species functional diversity within the PIN auxin efflux protein family. Elife. doi: 10.7554/eLife.31804

Open Access

Devin O’Connor and Ottoline Leyser (SLCU) lead this research that bridges the divide between a model dicot (Arabidopsis) and a model monocot (Brachypodium)as they investigate mechanisms of auxin transport, focussed on the PIN protein family. Arabidopsis lacks a clade of PIN proteins (termed Sister-of-PIN1 (SoPIN1) that are found in other plant species. They show that Brachypodium sopin1 mutants have inflorescence defects similar to Arabidopsis pin1 mutants, a similarity of function that is confirmed by the ability of soPIN1 to rescue the phenotype of null Atpin1 plants. However Brachy PIN1 is only able to rescue a less severe Atpin1 mutant. Overall they demonstrate that PIN1 functional specificity is determined by membrane and tissue-level accumulation and transport activity. As this paper is published in Elife, the journal provides reviewer comments and in this case they show that this manuscript was initially rejected. However the authors persisted and provided a reworked manuscript that convincing the reviewers that this study was appropriate for publication in Elife. An excellent lesson in persistence!


Gnan S, Marsh T, Kover PX (2017) Inflorescence photosynthetic contribution to fitness releases Arabidopsis thaliana plants from trade-off constraints on early flowering PLoS One doi: 10.1371/journal.pone.0185835

Open Access

In this study from Paula Kover’s lab at the University of Bath they investigate how the photosynthetic capacity of the Arabidopsis influoresence influences the time of flowering in a range of accessions. Interestingly after plants had flowering the authors removed rosette leaves to assess the ability of the influoresence to support future plant growth. Surprisingly there was a wide variation in general fitness following leaf removal, ranging from a growth reduction of 65% to no observed loss in fitness. These changes are due to both the differencies in the flowering time and in the number of lateral branches. This can explain how early flowering accessions can maintain fitness despite reduced vegetative growth.


Huang Z, Footitt S, Tang A, Finch-Savage WE (2017) Predicted global warming scenarios impact on the mother plant to alter seed dormancy and germination behavior in Arabidopsis Plant Cell Environ. doi: 10.1111/pce.13082

William Finch-Savage (University of Warwick) leads this investigation into the effect of temperature on seed development and dormancy. They used specially designed polyethylene tunnels that allowed in vivo variations in temperature and light conditions. Perhaps unsurprisingly they showed that temperature plays a significant role in future seed development with lower temperatures promoting dormancy but higher temperatures reduced dormancy that subsequently alters the timing of future life cycles, which has consequences for the species fitness.


Dzimitrowicz N, Breeze E, Frigerio L (2018) Long-Term Imaging of Endoplasmic Reticulum Morphology in Embryos During Seed Germination. Methods Mol Biol. doi: 10.1007/978-1-4939-7389-7_6

Lorenzo Frigerio (University of Warwick) leads this methods paper that describes the imaging of the endoplasmic reticulum over long periods during seed germination.


Ndinyanka Fabrice T, Kaech A, Barmettler G, Eichenberger C, Knox JP, Grossniklaus U, Ringli C (2017) Efficient preparation of Arabidopsis pollen tubes for ultrastructural analysis using chemical and cryo-fixation. BMC Plant Biol. doi: 10.1186/s12870-017-1136-x

Paul Knox (University of Leeds) is a co-author on this methods paper that outlines the necessary steps for efficient preparation of pollen tubes for subsequent ultrastructural analysis.

Arabidopsis Research Roundup: October 5th

After a brief hiatus the UK Arabidopsis Research Roundup returns with eight papers that focus on different aspects of Arabidopsis cell biology.

Firstly GARNet PI Jim Murray leads a study that performs a genome-wide analysis of sub-nucleosomal particles whilst Phil Wigge’s lab at SLCU conducts a more focused study on G-box regulatory sequences.

Thirdly Veronica Grieneisen (JIC) and co-workers have modelled the process of boron transport in the root, revealing exciting insights into how traffic jams might form.

Fourthly is a large scale biology paper led by Miriam Gifford (University of Warwick) that looks at the temporal and spatial expression patterns that control lateral root development.

Next Alexander Ruban (QMUL) investigates how low-light acclimated plants respond to high light.

The sixth and seventh studies are led by Alison Baker (Leeds) or Bill Davies (Lancaster) and look at phosphate or hormone signaling respectively.

Finally Gareth Jenkins (University of Glasgow) compares the UV-B signaling module in lower plants with that in Arabidopsis.


Pass DA, Sornay E, Marchbank A, Crawford MR, Paszkiewicz K, Kent NA, Murray JAH (2017) Genome-wide chromatin mapping with size resolution reveals a dynamic sub-nucleosomal landscape in Arabidopsis. PLoS Genet. doi: 10.1371/journal.pgen.1006988

Open Access

GARNet PI Jim Murray is the corresponding author on this study that performs a whole-genome scan of sub-nucleosomal particles (subNSPs) that have been identified using differential micrococcal nuclease (MNase) digestion. They link the position of subNSPs with RNAseq data taken from plants grown in different light conditions. They show that this new technique is able to discriminate regulatory regions that have been obscured by previous experimental procedures and therefore represents a very useful experimental method.


Ezer D, Shepherd SJ, Brestovitsky A, Dickinson P, Cortijo S, Charoensawan V, Box MS, Biswas S, Jaeger K, Wigge PA (2017) The G-box transcriptional regulatory code in Arabidopsis. Plant Physiol. 10.1104/pp.17.01086

Open Access

Phil Wigge (SLCU) is the corresponding author of this study that investigates the sequence elements that are linked to the conserved G-box regulatory motifs. They identify a set of bZIP and bHLH transcription factors that predict the expression of genes downstream of perfect G-boxes. In addition they have developed a website that provide visualisations of the G-box regulatory network (araboxcis.org).


Sotta N, Duncan S, Tanaka M, Takafumi S, Marée AF, Fujiwara T, Grieneisen VA (2017) Rapid transporter regulation prevents substrate flow traffic jams in boron transport. Elife. doi: 10.7554/eLife.27038

Open Access

Veronica Grieneisen (JIC) is the lead author on this detailed analysis of the regulatory circuits that are established during boron uptake in Arabidopsis roots. They used mathematical modelling to show that during boron uptake, swift regulation of transport activity is needed to prevent toxic accumulation of the metal. This system has analogy to the way in which traffic jams of nutrient flow might form and has relevance for regulatory systems outside of plant science. https://www.sciencedaily.com/releases/2017/09/170905104358.htm


Walker L, Boddington C, Jenkins D, Wang Y, Grønlund JT, Hulsmans J, Kumar S, Patel D, Moore JD, Carter A, Samavedam S, Bomono G, Hersh DS, Coruzzi GM, Burroughs NJ, Gifford ML (2017) Root architecture shaping by the environment is orchestrated by dynamic gene expression in space and time. Plant Cell. doi: 10.1105/tpc.16.00961

Open Access

Miriam Gifford (University of Warwick) leads this broad consortium that has taken a systems biology approach to better define the environmental factors that control dynamic root architecture. They track transcriptional responses during lateral root development in remarkable detail, looking at individual transcripts. They confirm the idea that the activity of a gene is not simply a function of its amino acid sequence but rather the temporal and spatial regulation of its expression.


Tian Y, Sacharz J, Ware MA, Zhang H, Ruban AV (2017) Effects of periodic photoinhibitory light exposure on physiology and productivity of Arabidopsis plants grown under low light. J Exp Bot. doi: 10.1093/jxb/erx213. Open Access

Alexander Ruban (QMUL) is the corresponding author on this collaboration with Chinese colleagues that examined the effect of high-light stress on low-light acclimated Arabidopsis plants. Initially these plants showed significant photo-inhibition but that they recovered rapidly and after 2 weeks of treatment there was no change in photosynthetic yield. In addition high light acclimated plants showed accelerated reproductive phase change that coincided with higher seed yield.


Qi W, Manfield IW, Muench SP, Baker A (2017) AtSPX1 affects the AtPHR1 -DNA binding equilibrium by binding monomeric AtPHR1 in solution. Biochem J. doi: 10.1042/BCJ20170522 Open Access

Alison Baker (University of Leeds) leads this research that focusses on the binding of the Phosphate Starvation Response 1 (PHR1) transcription factor to regulatory P1BS DNA sequences. They show a tandem P1BS sequence is bound more strongly than a single P1BS site. Ultimately they demonstrate tight regulation of phosphate signaling both by the concentration of phosphate as well as the activity of the interacting SPX protein.


Li X, Chen L, Forde BG, Davies WJ (2017) The Biphasic Root Growth Response to Abscisic Acid in Arabidopsis Involves Interaction with Ethylene and Auxin Signalling Pathways. Front Plant Sci. doi: 10.3389/fpls.2017.01493 Open Access

Bill Davies and Brian Forde (Lancaster University) lead this work that investigates the effect on ethylene and auxin on the biphasic response to ABA during root elongation. They used a range of hormone signalling mutants to show that the response to high ABA is via both ethylene and auzin signalling. In contrast the response to low ABA does not require ethylene signalling.


Soriano G, Cloix C, Heilmann M, Núñez-Olivera E, Martínez-Abaigar J, Jenkins GI (2017) Evolutionary conservation of structure and function of the UVR8 photoreceptor from the liverwort Marchantia polymorpha and the moss Physcomitrella patens. New Phytol. doi: 10.1111/nph.14767

Gareth Jenkins (University of Glasgow) is the corresponding author of this work that looks at the role of the UVR8 UV-B receptor in lower plants. They expressed the versions of UVR8 from a moss or a liverwort in Arabidopsis and showed that although there appears to be differences in the regulation of this protein, the mechanism of UV-B signaling is evolutionarily conserved

Arabidopsis Research Roundup: August 23rd

There is a bumper crop of papers in this weeks UK Arabidopsis Research Roundup! First up is a remarkable piece of work from George Bassel’s (University of Birmingham) lab that defines the network of cellular interactions that occur in the hypocotyl. Second and third are papers from the JIC in which Lars Ostergaard’s group uncovers the extent of the ETTIN signaling network and Caroline Dean‘s and Martin Howard’s labs provide evidence for a two step progression toward stable gene silencing following vernalisation at the FLC locus. Fourthly is a study that includes members of Alex Webb’s group (University of Cambridge) as co-authors that investigates the link between the circadian clock and night time starch metabolism. Fifth is a paper from Christine Foyer (University of Leeds) that looks at the effect of commonly used inhibitors on cellular redox state and gene expression. The next paper includes Phillip Carella (SLCU) as a co-author and looks at the role of classic flowering time genes on the phenomenon of Age-Related Resistance and finally Lee Sweetlove’s (University of Oxford) lab has published a methods paper for the analysis of photorespiration in non-photosynthetic tissues.


Jackson MD, Xu H, Duran-Nebreda S, Stamm P, Bassel GW (2017) Topological analysis of multicellular complexity in the plant hypocotyl. Elife http:/​/​dx.​doi.​org/10.7554/eLife.26023

Open Access

George Bassel (University of Birmingham) is the corresponding author on this work that provides fantastic images of the plant hypocotyl taken as part of an analysis on the cell growth dynamics in this organ. They show that Arabidopsis epidermal atrichoblast cells demonstrate a reduced path length that coincides with preferential movement of small molecules through these cells. They analysis this process in various mutants showing which gene activities were necessary for the construction of this pattern. In addition they compared topological features in Arabidopsis, Poppy and Foxglove, showing that cell interactions and path length determinants differ between these organisms. Overall this manuscript defines the network principles that control complex organ construction as well as a function for higher order patterning.


Simonini S, Bencivenga S, Trick M, Ostergaard L (2017) Auxin-Induced Modulation of ETTIN Activity Orchestrates Gene Expression in Arabidopsis. Plant Cell 10.1105/tpc.17.00389

Open Access

Last year Lars Ostergaard (JIC) discussed a paper from his lab on the GARNet YouTube channel in which they defined a new auxin-signaling paradigm that involved the non-canoical Auxin Response Factor ETTIN. This follow up to that study investigates the genetic network controlled by ETTIN activity and defines a range of developmental processes dependent on ETTIN auxin sensing. Furthermore by looking at direct ETTIN targets they suggest that this protein acts as a central node for the coordination of auxin signaling in the shoot. Finally their analysis of the effect of auxin on interactions between ETTIN and other transcription factors indicates that these are important factors in the diverse set of growth process controlled by auxin.


Yang H, Berry S, Olsson TSG, Hartley M, Howard M, Dean C (2017) Distinct phases of Polycomb silencing to hold epigenetic memory of cold in Arabidopsis. Science 10.1126/science.aan1121

This is another manuscript resulting from the extremely fruitful collaboration between the labs of Caroline Dean and Martin Howard at the John Innes Centre. This paper again focuses on the FLC locus and provides evidence for a new mechanism that defines how the binding of a subset of PRC2 factors nucleates a small region (<500bp) of chromatin at the FLC TSS, causing an increase in the repressive H3K27me2 histone mark. This metastable region serves as the seed for the development of stable epigenetic marks across the length of the locus through the activity of other distinct Polycomb factors. This occurs after a cold treatment and causes the spread of H3K27me2 repression. The novelty of this work is in the distinct temporal separation of phases of silencing, which ultimately result in the repression of FLC expression after a prolonged cold treatment.


Seki M, Ohara T, Hearn TJ, Frank A, da Silva VCH, Caldana C, Webb AAR, Satake A (2017) Adjustment of the Arabidopsis circadian oscillator by sugar signalling dictates the regulation of starch metabolism. Sci Rep. 10.1038/s41598-017-08325-y

Open Access

Research from Alex Webb’s group at the University of Cambridge features in the ARR for the second consecutive week, again on a similar topic. On this occasion they collaborate with Japanese colleagues to investigate the role of the circadian clock on determining the nighttime usage rate of starch that has accumulated during the day. They used a phase oscillator model to explain the link between the speed of the clock, starch breakdown and the maintenance of sucrose homeostasis. In addition they use Arabidopsis sugar response mutants to show that the circadian clock measures amount of cellular sucrose, which then controls the dynamics of starch breakdown.


Karpinska B, Alomrani SO, Foyer CH (2017) Inhibitor-induced oxidation of the nucleus and cytosol in Arabidopsis thaliana: implications for organelle to nucleus retrograde signalling. Philos Trans R Soc Lond B Biol Sci. 10.1098/rstb.2016.0392 Open Access

Christine Foyer (University of Leeds) is the corresponding author on this paper that looks at the effect of cellular oxidation on retrograde signaling between chloroplasts, mitochondria and the nucleus. They use a novel in vivo redox reporter to measure the effect of commonly used organelle inhibitors on cellular redox state. They discovered that these inhibitors cause a variety of effects on redox state and gene expression, which differed dependent on cell type. Researchers should be aware of these effects when they are drawing conclusions from their own experiments using these inhibitors.


Wilson DC, Kempthorne CJ, Carella P, Liscombe DK, Cameron R (2017) Age-Related Resistance in Arabidopsis thaliana Involves the MADS-domain Transcription Factor SHORT VEGETATIVE PHASE and Direct Action of Salicylic Acid on Pseudomonas syringae. Mol Plant Microbe Interact 10.1094/MPMI-07-17-0172-R

Phillip Carella is a Research Fellow at SLCU and this work from this previous lab in Canada investigates Arabidopsis Age-Related Resistance (ARR), a process that requires SA accumulation, which is then thought to act as an antimicrobial agent. The ARR response is lacking in plants containing a mutation in for the SHORT VEGETATIVE PHASE (SVP) gene. These svp plants have reduced SA, thought to be due to uncoupled overactivity of the SUPPRESSOR OF OVEREXPRESSION OF CO 1 gene. Overall this study shows that the flowering time gene SVP plays a complementary role in the control of SA accumulation, which confers ARR to older plants.


Fernie AR, Bauwe H, Sweetlove LJ (2017) Investigating the Role of the Photorespiratory Pathway in Non-photosynthetic Tissues. Methods Mol Biol 10.1007/978-1-4939-7225-8_15

Lee Sweetlove (University of Oxford) describes a protocol for evaluating the role of the photorespiration on the control of growth in non-photosynthetic tissues. This can be scaled for use in both Arabidopsis and in larger plants.

Arabidopsis Research Roundup: May 17th

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Published on: May 17, 2017

This weeks Arabidopsis research roundup begins with a paper led by CPIB at the University of Nottingham that discovers a somewhat surprisingly mechanism controling Arabidopsis root hydrotropism. Next are two papers from the University of Leeds that firstly investigate how the JAGGED LATERAL ORGANS gene influences the auxin response and secondly looks at the role of redox regulation in the control of the cell cycle and seed development. Finally are two papers that look at different aspects of the plant pathogen interactions. Jonathan Jones from the John Innes Centre is a co-author on a paper that dissects the multiple gene expression networks that control plant immunity whilst Charles Melnyk at the Sainsbury lab in Cambridge is involved with work that investigates the hormonal control mechanisms that influence the invasion of parasitic plants.


Dietrich D, Pang L, Kobayashi A, Fozard JA, Boudolf V, Bhosale R, Antoni R, Nguyen T, Hiratsuka S, Fujii N, Miyazawa Y, Bae TW, Wells DM,, Owen MR,, Band LR,, Dyson RJ, Jensen OE, King JR, Tracy SR, Sturrock CJ,, Mooney SJ, Roberts JA, Bhalerao RP, Dinneny JR, Rodriguez PL, Nagatani A, Hosokawa Y, Baskin TI, Pridmore TP, De Veylder L, Takahashi H, Bennett MJ (2017) Root hydrotropism is controlled via a cortex-specific growth mechanism. Nature Plants

http:/​/​dx.​doi.​org/10.1038/nplants.2017.57

Open Access via access link: http://rdcu.be/rSsk

Malcolm Bennett (University of Nottingham) leads a broad international collaboration that looks at the response of Arabidopsis roots to water. Surprisingly they show that this response occurs not in the root meristem but in the elongation zone and is controlled by a ABA signaling mechanism. They show that hydrotropism is dependent on cell elongation in the cortex but not in any other cell file. This is different to the gravitropic response and demonstrates that these tropisms are controlled by distinct tissue-specific mechanisms. To provide for information about this paper, lead author Daniela Dietrich joins Professor Bennett to discuss this paper on the GARNet YouTube channel and speculate on the function of different root cell layers in water uptake.


Rast-Somssich MI, Žádníková P, Schmid S, Kieffer M, Kepinski S, Simon R (2017) The Arabidopsis JAGGED LATERAL ORGANS (JLO) gene sensitizes plants to auxin. J Exp Bot.

http:/​/​dx.​doi.​org/10.1093/jxb/erx131 Open Access

This German-led study includes Stefan Kepinski (University of Leeds) as a co-author. They investigate the role of the JAGGED LATERAL ORGANS (JLO) transcription factor in the establishment of the stem cell niche in the root meristem. JLO interacts with auxin signaling pathway by influencing the degradation of the key regulator BODENLOS (BDL) via the TIR1-mediated degradation pathway. In jlo mutants BDL remains present in the meristem, which does not correctly develop. They discover a novel regulatory mechanism wherein the dosage of the TIR1 and AFB1 auxin receptors is reduced, which in turn prevents BDL degradation. This shows that the JLO transcription factor is a key upstream regulator of meristem formation by playing a significant role in the fine control of the auxin response.


De Simone A, Hubbard R, Vinegra de la Torre N, Velappan Y, Wilson M, Considine MJ, Soppe W, Foyer CH (2017) Redox changes during the cell cycle in the embryonic root meristem of Arabidopsis thaliana. Antioxid Redox Signal. http:/​/​dx.​doi.​org/10.1089/ars.2016.6959

Open Access

Christine Foyer (University of Leeds) collaborates with Australian and German researchers to unpick the role that redox regulation plays in the control of the plant cell cycle. They use an in vivo redox reporter (roGFP2) to show that a cycle of reduction and oxidation occurs throughout the cell cycle. Their experimental system is Arabidopsis seed germination and they show that vitamin c defective mutants with low redox buffering capacity have altered germination rates that coincide with a changed dry seed transcriptome. Overall this paper demonstrates that the cell cycle and embryo size are linked to redox regulation.


Hillmer RA, Tsuda K, Rallapalli G, Asai S, Truman W, Papke MD, Sakakibara H, Jones JDG, Myers CL, Katagiri F (2017) The highly buffered Arabidopsis immune signaling network conceals the functions of its components. PLoS Genet. http:/​/​dx.​doi.​org/10.1371/journal.pgen.1006639

Open Access

Jonathan Jones (John Innes Centre) is a co-author on this Japanese-led research that studies the signaling networks invovled in plant immunity. They use a systems biology approach to dissect the network of interactions that occur within the transcriptome when plants are exposed to the immune stimulant flagellin-22. This analysis discovers that there are separated networks that represent pathways controlled by different higher-level signals, such as jasmonate or salicylic acid. This provides the entire network with a degree of buffering that allows a more effective response to pathogen attack. This type of network analysis is able to reveal facets of the defence response that would not be possible when using simple null mutant analysis so adds consideration detail to the already complicated story of plant-pathogen interactions


Spallek T, Melnyk CW, Wakatake T, Zhang J, Sakamoto Y, Kiba T, Yoshida S, Matsunaga S, Sakakibara H, Shirasu K (2017) Interspecies hormonal control of host root morphology by parasitic plants. PNAS

http:/​/​dx.​doi.​org/10.1073/pnas.1619078114

Charles Melnyk (Sainsbury Lab, Cambridge) is an author on this study led by Ken Shirasu at RIKEN and uses Arabidopsis to investigate the relationship between parasitic plants and their hosts, specifically at the level of interspecies transport via a structure called the haustorium. Haustoria are structures through which substances, such as RNA and proteins, reciprocally move between host and parasite. In this paper they look at the interaction between Arabidopsis roots and the hemiparasitic plant Phtheirospermum japonicum, demonstrating that movement of molecules between species occurs via haustoria once a vascular connection is made. Arabidopsis secondary root growth is induced under infection, a response that requires the effect of the hormone cytokinin. They look at the genetics of this interaction and show that cytokinin signaling genes are important in establishing root hypertrophy. Overall this study demonstrates the important of cytokinin during infection with parasitic plants and might be an important target to design strategies to combat these negative interactions in systems.

Arabidopsis Research Roundup: March 17th

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Published on: March 17, 2017

This weeks UK Arabidopsis Research Roundup includes three papers featuring researchers from the University of Nottingham as well as manuscripts from Leeds, Lancaster, QMUL and The Sainsburys Lab in Norwich

Firstly Stefan Kepinski (Leeds) leads a study that investigates how Gravitropic Set Point Angle (GSA) is controlled in response to different growth factors. Secondly are two Methods papers featuring researchers from CPIB in Nottingham, the first of which is in collaboration with Lancaster University and introduces the Microphentron, which is an automated phenotyping platform that can be used for chemical biology screens. The second paper describes a non-destructive method for imaging floral tissues using CT scanning.

Ranjan Swarup is also a member of CPIB and in the next paper he has collaborated with French colleagues to investigate the role of SHR on root development in rice.

The fourth paper includes Cyril Zipfel as a co-author and investigates the role of damage-associated molecular patterns (DAMPs) in the response to pathogen attack whereas this weeks final paper is from the lab of Alexander Ruban (QMUL) and discovers the phenotypic consequences of persistent damage to PSII by photoinhibition.


Suruchi Roychoudhry, Martin Kieffer, Marta Del Bianco, Che-Yang Liao, Dolf Weijers Stefan Kepinski (2017) The developmental and environmental regulation of gravitropic setpoint angle in Arabidopsis and bean Scientific Reports http://dx.doi.org/10.1038/srep42664

Open Access

Stefan Kepinski (University of Leeds) leads this study that involves a collaboration with Dolf Weijers from Wageningen University. They investigate the role of both auxin and environmental factors in determining gravitropic set point angle (GSA), which is a measure of the growth of lateral organs away from primary shoots and roots. They show that nitrogen and phosphorous deficiency causes opposing effects on lateral root GSA, each of which are auxin-dependent. This contrasts with previous findings from work using bean adventitious roots. They find that these differences are maintained when Arabidopsis and bean roots are treated with different auxin concentrations. Latterly they also look at the effect of different light conditions on shoot GSA and put these findings into the context of potentially altering crop growth.

Stefan takes some time to discuss this paper for the GARNet YouTube Channel.


Burrell T, Fozard S, Holroyd GH, French AP, Pound MP, Bigley CJ, James Taylor C, Forde BG (2017) The Microphenotron: a robotic miniaturized plant phenotyping platform with diverse applications in chemical biology. Plant Methods

http://dx.doi.org/10.1186/s13007-017-0158-6 Open Access

This methods paper is a collaboration between the Universities of Lancaster and Nottingham led by Brian Forde that describes the Microphenotron. This device has been developed to facilitate chemical biology screens on in vivo plant tissues. This allows for the automated screening of either dicot or monocot roots or aerial tissues that have been grown on media infused with whichever chemical is relevant for the intended expriments. In situ GUS screening is also possible allowing for researchers to integrate information about growth and gene expression. The use of ‘Phytostrips’ in a 96-well format allows for high-throughput screening that is aligned with AutoRoot automated image analysis software to provide a rapid and facile method for undertaking small scale phenotypic screens. The Microphenotron facility is housed at the Lancester University, who are extremely open to collaboration so please get in contact if you are interested in using the facility.


Tracy SR, Gómez JF, Sturrock CJ, Wilson ZA, Ferguson AC (2017) Non-destructive determination of floral staging in cereals using X-ray micro computed tomography (µCT) Plant Methods. http://dx.doi.org/10.1186/s13007-017-0162-x Open Access

Alison Ferguson is the corresponding author on this methods paper that includes GARNet committee member Zoe Wilson and Saoirse Tracy from Dublin. They have developed a technique using X-ray µCT scanning to image developing flowers in Arabidopsis and barley plants, taking advantage of the excellent Hounsfield facility at the University of Nottingham. They show that the technique can be hugely beneficial for plant phenotyping by providing a non-destructive method of analyzing live floral development and how this can response to changes in the growth environment. Members of the Hounsfield facility are happy to discuss any potential collaborative work and future access to these type of facilities will hopefully be improved through the UKs involvement in the pan-european EMPHASIS project.


Henry S, Dievart A, Fanchon D, Pauluzzi G, Meynard D, Swarup R, Wu S, Lee CM, Gallagher K, Périn C (2017) SHR overexpression induces the formation of supernumerary cell layers with cortex cell identity in rice. Dev Biol. http://dx.doi.org/10.1016/j.ydbio.2017.03.001

Ranjan Swarup (CPIB) is a co-author on this study that includes French and US researchers. Previously they had shown that expression of rice SHORTROOT (OsSHR) genes could compliment the Arabidopsis shr mutant. In this study they show that overexpression of OsSHR and AtSHR in rice roots causes growth of wider, shorter roots that have an increased number of cortical cell layers. This demonstrates that the mechanisms that control the differentiation of cortical cell layers is conserved throughout land plants, with SHR being a key determinant in this process.


de Azevedo Souza C, Li S, Lin AZ, Boutrot F, Grossmann G, Zipfel C, Somerville S (2017) Cellulose-derived oligomers act as damage-associated molecular patterns and trigger defense-like responses. Plant Physiol. http://dx.doi.org/10.1104/pp.16.01680

Cyril Zipfel (The Sainsbury Lab) is a co-author on this study from the lab of Shauna Somerville in California that focuses on the concept of damage-associated molecular patterns (DAMPs). These can be defined as cell wall breakdown components and stimulate the same defence responses as more fully characterised pathogen- or microbe-associated molecular patterns (PAMPs). Intuitively this makes sense as during infection many pathogens will cause cell wall breakdown. The authors show that cellulose-derived oligomers trigger a signalling response similar to that caused by oligogalacturonides or chito-oligomers but that lacks an increase in ROS or in callose deposition. These results confirm that cellulose-derived signals feed into the plants mechanism for cell wall scanning and acts synergistically with other signals that result from pathogen attack.


Tian Y, Ungerer P, Zhang H, Ruban AV (2017) Direct impact of the sustained decline in the photosystem II efficiency upon plant productivity at different developmental stages. J Plant Physiol. http://dx.doi.org/110.1016/j.jplph.2016.10.017

Image from http://www.sciencedirect.com/science/article/pii/S0176161717300433

Alexander Ruban (QMUL) leads this Sino-UK collaboration that investigates how the photoinhibiton of photosystem II impacts overall plant growth. In this study they use lincomycin to block chloroplast protein synthesis, which prevents the plant from restoring PSII function after photoinhibitory damage. Treated plants accumulate less starch and showed reduced above-ground biomass. This leads to a decrease in seed yield. Perhaps unsurprisingly this research shows that restoring the full function of PSII after photoinhibition to key to maintaining normally functioning electron transport rate that leads into metabolic production and growth rate.

Arabidopsis Research Roundup: July 19th

There are six papers in this weeks Arabidopsis Research Roundup. Two of these include research on the stomatal patterning gene TMM. Firstly a White Rose consortium investigates the ancestral basis of stomatal patterning, whilst a Glasgow-based study investigates the relationship between patterning and the dynamics of guard cell opening. The GARNet committee is represented by work from Cardiff that looks at the relationship between seed size and shoot branching and also from Cambridge in research that studies meiotic recombination in genomic regions important for pathogen defense. Finally are two studies that look into aspects of root and shoot patterning and include co-authors from CPIB in Nottingham or the John Innes Centre.

Caine R, Chater CC, Kamisugi Y, Cuming AC, Beerling DJ, Gray JE, Fleming AJ (2016) An ancestral stomatal patterning module revealed in the non-vascular land plant Physcomitrella patens Development

http://dx.doi.org/10.1242/dev.135038 Open Access

This study is a collaboration between labs in Sheffield and Leeds, led by Andrew Fleming (Sheffield). They investigate the role that the signalling module comprised of Epidermal Patterning Factors (EPFs), ERECTA and TMM play during the evolution of stomatal patterning. This module is known to play an important role in Arabidopsis and in this study the authors show that the moss Physcomitrella patens contains homologs of each of the genes and that they perform the same function. When P.paten versions of these genes are transferred to equivalent Arabidopsis mutants they show conserved function demonstrating that this module is an example of an ancestral patterning system.

Andrew Fleming provides a brief audio description of this manuscript:

Papanatsiou M, Amtmann A, Blatt MR (2016) Stomatal spacing facilitates guard cell ion transport independent of the epidermal solute reservoir. Plant Physiol. http://dx.doi.org/10.1104/pp.16.00850 Open Access

Mike Blatt and Anna Amtmann (University of Glasgow) are the co-supervisors for this study into the relationshop between ion transport in stomatal guard cells and their physical positioning within a leaf. They used a genetic approach to assess the effect of stomatal clustering, showing that too many mouths (tmm) mutant plants have reduced stomatal movements associated with alterations in K+ channel gating and coincident with a surprising reduction in the level of K+ ions in guard cells. These results underline the importance of stomatal spacing in this process but do not provide a full explanation into the alteration in K+ ion dynamics.

Sornay E, Dewitte W, Murray JAH (2016) Seed size plasticity in response to embryonic lethality conferred by ectopic CYCD activation is dependent on plant architecture Plant Signaling and Behaviour e1192741

http://dx.doi.org/10.1080/15592324.2016.1192741 Open Access

From http://dx.doi.org/10.1080/15592324.2016.1192741
From http://dx.doi.org/10.1080/15592324.2016.1192741

This research comes from the lab of GARNet PI Jim Murray (Cardiff) and investigates cell proliferation and growth within a developing seed. They previously have shown that targeting of D-type cyclin CYCD7;1 to the central cell and early endosperm can trigger nuclear divisions and ovule abortion, which leads to a smaller number of larger seed. In this study they show that development of larger seed in transgenic plants is influenced by the architecture of the mother, as plants with increased side branches, caused by pruning of the main stem, do not generate this phenotype. This is indicative of a close relationship between the amount of resources allocated to different parts of the plant and that a transgenic effect was altered by a different plant morphology. This should provide an important insight into future work that aims to define the effect of any particular transgenic alteration.

Choi K, Reinhard C, Serra H, Ziolkowski PA,, Underwood CJ,, Zhao X, Hardcastle TJ, Yelina NE, Griffin C, Jackson M, Mézard C, McVean G, Copenhaver GP,, Henderson IR (2016) Recombination Rate Heterogeneity within Arabidopsis Disease Resistance Genes. PLoS Genet. 12(7):e1006179.

http://dx.doi.org/10.1371/journal.pgen.1006179 Open Access

GARNet advisory board member Ian Henderson (Cambridge) is the corresponding author of this study that involves contributions from the UK, US, Poland and France. They investigate genomic regions that show increased meiotic recombination, which is predicted to occur coincident with genes involved in pathogen defence given their requirement to adapt to new external challenges. This study focuses on NBS-LRR domain proteins that tend to physically cluster in the Arabidopsis genome. Interesting they discovered both hot and coldspots for meiotic recombination that associate with NBS-LRR clusters, the later often correlating with structural heterozygosity. In a more detailed dissection of 1000 crossovers in the RESISTANCE TO ALBUGO CANDIDA1 (RAC1) R hotspot, they discovered higher recombination frequencies associating with known sequence motifs important for the pathogen response, which were influenced by ecotype-specific factors. Ultimately the authors note that there is a complex relationship between regions of meiotic recombination, structural heterozygosity and the evolutionary pressures that occurs with host-pathogen relationships.

Orman-Ligeza B, Parizot B, de Rycke R, Fernandez A, Himschoot E, Van Breusegem F, Bennett MJ, Périlleux C, Beeckman T, Draye X (2016) RBOH-mediated ROS production facilitates lateral root emergence in Arabidopsis. Development http://dx.doi.org/10.1242/dev.136465 Open Access

From http://dx.doi.org/10.1242/dev.136465
From http://dx.doi.org/10.1242/dev.136465

 Malcolm Bennett (CPIB) is the sole UK-based co-author on this study led by Belgian collaborators and investigates the role of reactive oxygen species (ROS) in auxin-regulated lateral root (LR) formation. They show that ROS can reactivate LR primordia and pre-branch sites, resulting in increased LR numbers. This occurs in both wildtype and in auxin mutants that have reduced numbers due to changes in auxin-mediated cell wall remodeling. ROS is deposited in the apoplast of emerging LR cells in a pattern that is coincident with the expression of the RESPIRATORY BURST OXIDASE HOMOLOGS (RBOH) genes. Concomitantly the altered expression of RBOH was shown to affect the development and emergence of LRs. This adds a further level of complexity to the current understanding of the signaling factors that converge to facilitate LR growth.

 

Shi B,, Zhang C, Tian C, Wang J,, Wang Q,, Xu T,, Xu Y, Ohno C, Sablowski R, Heisler MG, Theres K, Wang Y, Jiao Y (2016) Two-Step Regulation of a Meristematic Cell Population Acting in Shoot Branching in Arabidopsis. PLoS Genet. http://dx.doi.org/10.1371/journal.pgen.1006168 Open Access

This Chinese-led study includes Robert Sablowski (JIC) as a co-author and studies the factors that influence the development of axillary meristems. They use innovative live imaging to show that SHOOT MERISTEMLESS (STM) is continuously expressed and that this dependent on a leaf axil auxin minimum. Once STM expression is lost then the axil is unable to form a meristem even if STM is switched back later in development, indicating that cells undergo an irreversible developmental commitment. The expression domain of STM is under cell-type specific control of REVOLUTA (REV) DNA binding. Overall this study demonstrates that meristematic competence and initiation is dependent on differing levels of the key regulator STM.

From http://dx.doi.org/10.1371/journal.pgen.1006168
From http://dx.doi.org/10.1371/journal.pgen.1006168

Arabidopsis Research Roundup: May 5th

There are a bumper crop of papers in this edition of the Arabidopsis Research Roundup. First from the University of Manchester is a paper that identifies a protein involved in plant programmed cell death. Secondly are two papers from the University of Bristol that highlight the role of viruses in the reflectivity of plant leaves and an assessment of the growth parameters of Arabidopsis on different soil-types. Thirdly are three papers from University of Edinburgh that either use CRISPR-Cas technology to develop virus-research plants, investigate the relationship between photoperiod and metabolism or present a method for assessment of protein S-nitrosylation. Fourthly is a paper that includes a contribution from the University of Leeds that investigates the evolutionary and functional relationship of the WOX gene family. Finally is a study that highlights the role of the AUGMIN complex during microtubule activity that includes a contribution from the University of Leicester.

In addition, although not involving Arabidopsis, we should mention an exciting study from Gerben van Ooijen (Edinburgh) that has discovered a conserved circadian mechanism based on magnesium rhythms that is linked to energy expenditure.

Ge Y, Cai YM, Bonneau L, Rotari V, Danon A, McKenzie EA, McLellan H, Mach L, Gallois P (2016) Inhibition of cathepsin B by caspase-3 inhibitors blocks programmed cell death in Arabidopsis. Cell Death Differ. http://dx.doi.org/10.1038/cdd.2016.34 Open Access

The corresponding author of this paper is Patrick Gallois from the University of Manchester and includes contributions from Hazel McLellan in Dundee almongside Chinese and Austrian collaborators. This study investigates the role of caspase proteins on control of programmed cell death in plants. This research area has been hindered by the apparent lack of plant caspase orthologues despite pharmacological evidence that proteins with caspase activity are active in plants. The authors use a labeled caspase-3 inhibitor to identify the Arabidopsis Cathepsin B3 (AtCathB3) protein as having caspase activity, which was verified using recombinant proteins during in vitro enzyme assays. AtCathepsinB1,2,3 triple mutant plants demonstrate a reduction in PCD induced by different stresses and explains why caspase inhibitors are effective tools for studying PCD in plants. The core Cathepsin B protein is evolutionarily conserved suggesting that an ancestral pathway exists that controls PCD, the details of which require further study.

Maxwell DJ, Partridge JC, Roberts NW, Boonham N, Foster GD (2016) The Effects of Plant Virus Infection on Polarization Reflection from Leaves. PLoS One. 11(4):e0152836 http://dx.doi.org/ 10.1371/journal.pone.0152836 Open Access

Gary Foster’s research group at the University of Bristol collaborate with others at the University of York and in Australia for this study that investigates how plant viruses may modify gene expression to benefit their own transmission. They show that Potato virus Y and Cucumber mosaic virus (CMW), which both are transmitted by aphids, significantly reduce the amount of polarised light that is reflected from abaxial leaf surfaces of tobacco plants particularly when compared to the effects caused by non-insect vectored viruses. However this effect was not shown in Arabidopsis leaves infected by a variety of differently transmitted viruses. Interestingly ECERIFERUM6 (CER6) transcripts accumulate to higher levels following infection with insect vectored viruses and as this gene is involved in cuticle wax synthesis the authors suggest that induced changes in cuticle composition might be key in understanding how viruses encourage predation by their insect vectors. Finally the authors discuss the overall adaptive significance of these results.

Drake T, Keating M, Summers R, Yochikawa A, Pitman T, Dodd AN (2016) The Cultivation of Arabidopsis for Experimental Research Using Commercially Available Peat-Based and Peat-Free Growing Media. PLoS One. 11(4):e0153625 http://dx.doi.org/ 10.1371/journal.pone.0153625 Open AccessPeatPic

GARNet committee member Anthony Dodd, also from the University of Bristol, leads this study into the growth of Arabidopsis on peat-free media, which was motivated by the unsustainable use of peat-based composts. They found that biomass accumulation and seed yield were reduced on peat-free media and that some types of this media was more suspectible to fungal contamination. Overall vegetative phenotypic parameters were similar between plants grown on peat-based or peat-free media, indicating that this type of media will be appropriate for future analysis. However the seed yield was usually reduced, indicating that experiments looking at post-phase change phenotypes might not be as comparable between plants growth on media with different amount of peat.

Pyott DE, Sheehan E, Molnar A (2016) Engineering of CRISPR/Cas9-mediated potyvirus resistance in transgene-free Arabidopsis plants Mol Plant Pathol. http://dx.doi.org/10.1111/mpp.12417

Attila Molnar (Edinburgh) is the corresponding author on this study that uses the transformative technology CRISPR/Cas9 to engineer Arabidopsis plants that are resistant to potyvirus infection. This is achieved by targeting the genes encoding the translation initiation factor eIF(iso)4E that had been previously identified as being critical for viral establishment. Importantly they subsequently selected transgene-free plants that have no phenotypic changes when compared to wildtype growth under standard conditions. As the potyvirus Turnip Mosaic Virus is an important pathogen for vegetable crops this is potentially an extremely powerful technique for generating virus-resistance food crops.

Flis A, Sulpice R, Seaton DD, Ivakov AA, Liput M, Abel C, Millar AJ, Stitt M (2016) Photoperiod-dependent changes in the phase of core clock transcripts and global transcriptional outputs at dawn and dusk in Arabidopsis Plant Cell Environ. http://dx.doi.org/10.1111/pce.12754

This German–led study aims to connect the expression of photoperiod-length responsive circadian clock-regulated genes with those involved in metabolic processes such as starch degradation and includes a contribution from Professor Andrew Miller from the Edinburgh SynthSys Centre. The authors assess global gene expression by transcript profiling at photoperiods ranging from 4-18 hours and found that changes in transcript abundance at dawn throughout these photoperiods were as large as changes seen in individual experiments when comparing dawn and dusk. These complex interactions revealed coordinated regulation of key metabolic processes and begins to demonstrate how metabolism is linked to photoperiod.

Homem RA, Le Bihan T, Yu M, Loake GJ (2016) Identification of S-Nitrosothiols by the Sequential Cysteine Blocking Technique Methods Mol Biol. 1424:163-74. http://dx.doi.org/10.1007/978-1-4939-3600-7_14

This paper from the lab of Gary Loake (Edinburgh) describes the methods they use to investigate the role of protein S-nitrosylation in the immune responses of Arabidopsis. These are based on a modification of the biotin-switch technique, which they term sequential cysteine blocking.

Dolzblasz A, Nardmann J, Clerici E, Causier B, van der Graaff E, Chen J, Davies B, Werr W, Laux T (2016) Stem cell regulation by Arabidopsis WOX genes Mol Plant. S1674-2052(16)30029-6 http://dx.doi.org/10.1016/j.molp.2016.04.007

This German-led study includes work from the lab of Brendan Davies at the University of Leeds and investigates the role of the WUSCHEL-RELATED HOMEOBOX (WOX) transcription factor gene family during stem cell development and maintenance. Most members of the WUS-clade can largely substitute for WUSCHEL activity in the shoot meristem, which is absolutely dependent on a conserved WUS-box motif that is critical for the interaction with TOPLESS co-repressors. In contrast to the WUS clade, the WOX13 and WOX9 clades cannot substitute for WUS activity. The indicates that WOX control of shoot and floral meristem relies on certain currently not-fully-understood attributes of the WUS-clade of proteins.

Oh SA, Jeon J, Park HJ, Grini PE, Twell D, Park SK (2016) Analysis of gemini pollen 3 mutant suggests a broad function of AUGMIN in microtubule organization during sexual reproduction in Arabidopsis Plant J. http://dx.doi.org/10.1111/tpj.13192

David Twell (Leicester) is an author on his Korean-led study that reports on the identification of the new gem3 mutant, which displays defects in gametophytic development. Mutant plants exhibits disrupted cell division during male meiosis, at pollen mitosis I and throughout female gametogenesis. Gem3 is a hypomorphic allele of the AUGMIN subunit 6 gene, which is a component of Augmin complex responsible for microtubule (MT) nucleation in acentrosomal cells. In the gem3 mutant, the authors show that MT arrays are incorrectly distributed, likely causing the gametophyte-specific phenotypes and demonstrating a broad role for the augmin complex during sexual reproduction in flowering plants

Arabidopsis Research Roundup: April 1st.

This weeks Arabidopsis Research Roundup contains an eclectic mix of investigations. Firstly is a study from Peter Unwin that investigates the molecular factors that control interactions between plants and nematode parasites. Secondly is a study led by John Christie that investigates the factors that control hypocotyl curvature. Thirdly is a fascinating proof-of-concept synbio-style study from Rothamstead Research where an algal gene is transferred into Arabidopsis in the hope of developing a phytomediation-based solution to heavy metal contamination. Fourthly is a study from David Bass that catalogues protist species that feed on leaf-microorganisms whilst finally John Carr heads a study that compares RNA-dependent RNA polymerases from Arabidopsis and Potato.

Eves-van den Akker S, Lilley CJ, Yusup HB, Jones JT, Urwin PE (2016) Functional C-terminally encoded plant peptide (CEP) hormone domains evolved de novo in the plant parasite Rotylenchulus reniformis. Mol Plant Pathol. http://dx.doi.org/10.1111/mpp.12402).CEP1

This study is a collaboration between researchers at the Universities of Dundee and Leeds, led by Peter Unwin. The focus of the paper is the interaction of Plant-Parasitic Nematodes (PPNs) with their plant hosts. PPNs stimulate redifferentiation of vascular tissues to form ‘feeding structures’ that benefit the parasite. This process is mediated by a diverse family of effector proteins termed C-terminally Encoded Peptide plant hormone mimics (CEPs). This study investigates the CEPs from the nematode Rotylenchulus reniformis and suggests that these peptides evolved de novo in this organism. They show that the activity of a synthetic peptide corresponding to RrCEPs causes a reduction in primary root elongation whilst upregulating a set of genes including the nitrate transporter AtNRT2.1. The authors propose that CEPs evolved in R. reniformis to allow sustained biotrophy by upregulating a specific set of feeding-responsive genes and by limiting the size of the feeding site produced. This study represents an exciting introduction to a currently under-researched area within plant-pathogen interactions.

Sullivan S, Hart JE, Rasch P, Walker CH, Christie JM (2016) Phytochrome A Mediates Blue-Light Enhancement of Second-Positive Phototropism in Arabidopsis. Front Plant Sci. 7:290 http://dx.doi.org/10.3389/fpls.2016.00290 Open AccessFrontiersPHOT1

John Christie (Glasgow) is the corresponding author on this investigation into the role of the blue-light receptor phototropin 1 (phot1) during hypocotyl phototropism. Curvature of this organ is enhanced by treatment by red-light mediated by the phytochromeA receptor. However this study shows that pre-treatment with blue-light can also enhance this hypocotyl curvature although this did not occur at higher light intensities. In addition phototropic enhancement was also lacking when PHOT1 is expressed only in the hypocotyl epidermis. Therefore the study shows that the phyA impact on phot1 signaling is restricted to low light intensities and in tissues other than the epidermis.

Zhong Tang, Yanling Lv, Fei Chen, Wenwen Zhang, Barry P. Rosen, and Fang-Jie Zhao (2016) Arsenic Methylation in Arabidopsis thaliana Expressing an Algal Arsenite Methyltransferase Gene Increases Arsenic Phytotoxicity J. Agric. Food Chem. http://dx.doi.org/10.1021/acs.jafc.6b00462 Open Access ArsM

This synthetic biology-focused study is led by Fang-Jie Zhao at Rothamstead Research. The authors take an algal gene (arsM) that allows the transformation of inorganic arsenic to a more volatile methylated version. The biological activity of this enzyme was successfully transferred to two different Arabidopsis ecotypes. However interestingly these transgenic plants became more sensitive to arsenic in growth media suggesting that the new methylated arsenic intermediate is more phytotoxic than inorganic arsenic. Therefore this study demonstrates a negative consequence of this project that attempted to engineer arsenic tolerance in plants. Once again this demonstrates that nature rarely acts predictably and any great ideas usually need to be tested in vivo.

Ploch S, Rose L, Bass D, Bonkowski M (2016) High Diversity Revealed in Leaf Associated Protists (Rhizaria: Cercozoa) of Brassicaceae J Eukaryot Microbiol. http://dx.doi.org/10.1111/jeu.12314

After a fantastic opening line in the abstract, ‘The largest biological surface on earth is formed by plant leaves’, this study includes the work of David Bass from the Natural History Museum in London. They investigate the abundance of protists that associate with leaf-inhabiting microorganisms, the “phyllosphere microbiome“. Their findings demonstrate that protists should be considered an important part of the diversity of plant-interacting microbial organisms.

Hunter LJ, Brockington SF, Murphy AM, Pate AE, Gruden K, MacFarlane SA, Palukaitis P, Carr JP (2016) RNA-dependent RNA polymerase 1 in potato (Solanum tuberosum) and its relationship to other plant RNA-dependent RNA polymerases Sci Rep. 6:23082 http://dx.doi.org/10.1038/srep23082 Open Access

John Carr (Cambridge) is the UK-lead on this collaboration with Slovenian and Korean researchers. They primarily investigate the role of the RDR1 RNA-dependent RNA polymerase (RDRs) in potato. In Arabidopsis the RDR1 gene contributes to basal viral resistance but potato plants deficient in StRDR1 do not show altered susceptibility to three different plant viruses. In addition they perform a phylogenetic analysis on the RDR genes and identify a novel RDR7 gene that is only found in Rosids (but not Arabidopsis.

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