Dr Mark Fricker

Associate Professor in Plant Sciences
Mark Fricker

Tel +44 (0) 1865 275015
Fax +44 (0) 1865 275074

Research Area

Imaging signalling and transport in complex systems

Research Description

Website: www.markfricker.org

Biological Network analysis

We have pioneered network analysis of foraging basidiomycete fungi growing in unconstrained microcosms (Bebber et al. 2007a; Bebber et al. 2007b; Boddy et al. 2009; Boddy et al. 2010; Fricker et al. 2008a; Fricker et al. 2007; Fricker et al. 2008b; Heaton et al. 2010; Rotheray et al. 2008). We have shown using graph-theoretic analysis of digitised networks, that these indeterminate, de-centralized systems can yield adaptive networks with both high transport capacity and robustness to damage, but at a relatively low cost, through a 'Darwinian' process of selective reinforcement of key transport pathways and recycling of redundant routes (Bebber et al. 2007a). Furthermore, fungal networks are able to dynamically modify link strengths and local connectivity when subject to experimental attack to readjust the balance between transport capacity, robustness to damage and resource allocation, resulting in increased resilience as the environment becomes more challenging (Boddy et al. 2010; Rotheray et al. 2008). Furthermore, genetically disrupting network formation in Neurospora reduces long distance nutrient transport (Simonin et al., 2012).

Currently network extraction is a laborious manual exercise so we have developed high-throughput, automated imaging, visualisation and network analysis protocols to extract biological network organisation using Phase Congruency Tensors (PCTs) to specifically enhance curvi-linear features (Obara, 2012a-c). This approach can rapidly extract networks with 105 links in a few minutes with high fidelity.

The underlying mechanisms leading to the emergence of adaptive behaviour in macroscopic mycelial networks are unknown. However, models based on growth-induced mass flow through the experimentally determined macroscopic networks provide a high level of explanatory power for the transport of added radiolabel (Heaton et al. 2010; Heaton et al. 2012). We infer that bio-physical hydraulic coupling and internal flows observed in macroscopic networks may act as the central mechanism enabling coordinated growth across the complete range of scales in networked organisms. These models can be extended to capture the energetic constraints that determine fungal life history strategy (Heaton et al. 2015).

Slime mold networks

The work on fungal networks has also led to collaboration with Toshiyuki Nakagaki and his team in Hokkaido to understand network formation in the acellular slime mold Physarum polycephalum. We have shown that it can form networks with comparable efficiency, fault tolerance, and cost to those of real-world infrastructure networks such as the Tokyo rail system (Tero et al. 2010; Kunita et al., 2014), which was awarded an IgNobel prize in 2010. The core mechanisms needed for adaptive network formation can be captured in a biologically inspired mathematical model that may be useful to guide network construction in other domains. Network formation in fungi and slime molds can also be compared to network architecture in other domains using mesoscale community detection and clustering algorithms (Onnela et al., 2012; Lee et al., 2015).



Long distance nutrient transport in fungi

Analysis of the network architecture allows predictive modeling of the expected nutrient dynamics. The pattern of nutrient distribution can be mapped in vivo using novel scintillation imaging techniques we have developed to map the transport of 14C-labelled α-amino isobutyrate (14C-AIB) as a non-metabolised, radiolabelled amino-acid analogue in Phanerochaete velutina (Tlalka et al. 2002, Tlalka et al. 2003, Bebber et al. 2007, Bebber et al. 2007, Fricker et al. 2008, Tlalka et al. 2008). We have revealed novel N-transport phenomena, including rapid, preferential N-resource allocation to C-rich sinks, induction of simultaneous bi-directional transport, and abrupt switching between different pre-existing transport routes (Tlalka et al., 2002; Tlalka et al., 2003; Bebber et al., 2007; Fricker et al., 2007; Tlalka et al., 2007). Using a simulation model we have also shown that preferential N-allocation and growth in response to a new resource in a range of resource environments confers an ecological benefit (Darrah et al., 2014). There is also a pulsatile component to transport and colonies self-organise into well demarcated domains that are identifiable by differences in the phase relationship of the pulses (Tlalka et al., 2003; Bebber et al., 2007; Fricker et al., 2007; Tlalka et al., 2007).

Imaging signal transduction

We have a long track record in the development and application of imaging techniques to map redox homeostasis and signalling plant and fungal systems (Moore et al., 2006; Brandizzi et al,. 2002). We developed a system for quantitative imaging of total cytoplasmic glutathione (GSH) in vivo following GST-catalysed conjugation to monochlorobimane (MCB) to give a fluorescent glutathione-bimane (GSB) adduct. This has allowed measurement of concentrations of GSH in defined cell types in intact tissues, to dissect the control of GSH synthesis and to analyse activities of GSH-based xenobiotic detoxification pathways in intact tissues with sub-cellular resolution. The imaging assay is technically complex (Fricker et al., 2000; Meyer and Fricker, 2000; Meyer et al., 2001), but now successfully applied to GSH measurements in several different types of tissues including roots (Sanchez-Fernandez et al., 1997; Fricker et al., 2001), trichomes (Gutiérrez-Alcalá et al 2000), suspension culture cells (Meyer and Fricker, 2002), and mesophyll, epidermal and guard cells of wild type and transgenic poplar over-expressing g-ECS (Hartmann et al 2003), mutants in ER morphology (Au et al., 2012), and most recently pathogenic fungi (Samalova et al., 2014).

This work on total GSH measurements has now been combined with in vivo redox imaging using ratiometric roGFP1 and roGFP2 (Schwarzlander et al., 2008; Marty et al., 2009) and application of these probes to measure mitochondrial redox state (Wagner et al., 2015) using custom ratio imaging software (Fricker, 2015) under a range of stress conditions (Lehmann et al., 2008; Schwarzlander et al., 2009), in mutants deficient in MnSOD (Morgan et al 2008), the mitochondrial Ca2+–channel subunit MICU1 (Wagner et al.,  2015) and during pathogen attack (Fuchs et al., 2015).

We have also shown that cpYFP does not act as a reporter for superoxide, but behaves as an  effective pH indicator in vivo, and reveals the presence of transient alkalinisation events associated with flickering in mitochondrial membrane potential (Schwarzlander et al. 2011; 2012).

We have started to move these approaches across to the pathogenic rice blast fungus, Magnaporthe oryzae, coupling the redox and ROS probes with nitric oxide measurements (Samalova et al., 2013; 2014).



Fuchs, R. Kopischke, M., Klapprodt, C. Hause, G., Meyer, A., Schwarzländer, M. Fricker, M.D. and Lipka, V. (2015). Immobilized subpopulations of leaf epidermal mitochondria mediate PEN2-dependent pathogen entry control in Arabidopsis. Plant Cell, (on line)


Lee, S.H., Fricker, M.D. and Porter, M.A. (2015) Mesoscale analyses of fungal networks as an approach to quantifying phenotypic traits. Network Science (in press). 

Wagner, S., Behera, S., De Bortoli, S., Logan, D., Fuchs, P., Carraretto, L., Teardo, E., Cendron, L., Nietzel, T., Füßl, M., Doccula, F., Navazio, L., Fricker, M., Van Aken, O., Finkemeier, I., Meyer, A., Szabo, I., Costa, A., and Schwarzländer, M. (2015) AtMICU choreographs mitochondrial Ca2+ dynamics in Arabidopsis. Plant Cell 27, 3190-3212.

Doi: 10.1105/tpc.15.00509 pdf

Heaton, L.L.M., Jones, N.S. and Fricker, M.D. (2015) Energetic constraints on saprotrophic fungal growth determine life history strategies. American Naturalist (in press). Doi: 10.1086/684392 pdf

Fricker, M.D. (2015) Quantitative Redox Imaging Software. Anti-oxidant and Redox Signalling (in press).

Doi: http://dx.doi.org/10.1089/ars.2015.6390 pdf

Wagner, S., Nietzel, T., Aller, I., Costa, A., Fricker, M.D., Meyer, A.J. and Schwarzländer, M. (2015) Analysis of plant mitochondrial function using fluorescent protein sensors. Methods in Molecular Biology 1305, 241-252.

Doi: https://doi.org/10.1007/978-1-4939-2639-8_17 pdf



Schwarzländer, M., Wagner, S., Ermakova, Y. G., Belousov, V. V., Radi, R., Beckman, J. S., Murphy, M. P. (2014). The 'mitoflash' probe cpYFP does not respond to superoxide. Nature, 514(7523), E12-E14.

Doi:10.1038/nature13858 pdf

Lichius, A., Goryachev, A. B., Fricker, M. D., Obara, B., Castro-Longoria, E., & Read, N. D. (2014). CDC-42 and RAC-1 regulate opposite chemotropisms in Neurospora crassa. J. Cell Science, 127, 1953-1965.

Doi:10.1242/jcs.141630 pdf

Littlejohn, G. R., Mansfield, J. C., Christmas, J. T., Witterick, E., Fricker, M. D., Grant, M. R., Love, J. (2014). An update: improvements in imaging perfluorocarbon-mounted plant leaves with implications for studies of plant pathology, physiology, development and cell biology. Frontiers in Plant Science, 5, 140.

Doi:10.3389/fpls.2014.00140 pdf

Samalova, M., Meyer, A. J., Gurr, S. J., & Fricker, M. D. (2014). Robust anti-oxidant defences in the rice blast fungus Magnaporthe oryzae confer tolerance to the host oxidative burst. New Phytol, 201, 556-573.

Doi:10.1111/nph.12530 pdf

Darrah, P. R., & Fricker, M. D. (2014). Foraging by a wood-decomposing fungus is ecologically adaptive. Environ. Microbiol., 16, 118-129.

Doi:10.1111/1462-2920.12216 pdf



Samalova, M., Johnson, J., Illes, M., Kelly, S., Fricker, M., & Gurr, S. (2013). Nitric oxide generated by the rice blast fungus Magnaporthe oryzae drives plant infection. New Phytol, 197, 207-222.

Doi:10.1111/j.1469-8137.2012.04368.x pdf

Kunita, I., Yoshihara, K., Tero, A., Ito, K., Lee, C.-F., Fricker, M.D. and Nakagaki, T. (2013) Adaptive path-finding and transport network formation by the amoeba-like organism Physarum. In Proceedings in Information and Communications Technology, 6, 14-29.

Doi: 10.1007/978-4-431-54394-7_2 pdf



Simonin, A., Palma-Guerrero, J., Fricker, M., & Louise Glass, N. (2012). Physiological significance of network organization in fungi. Eukaryotic Cell, 11, 1345-1352.

Doi:10.1128/EC.00213-12 pdf

Yasumura, Y., Pierik, R., Fricker, M. D., Voesenek, L. A., & Harberd, N. P. (2012). Studies of Physcomitrella patens reveal that ethylene-mediated submergence responses arose relatively early in land-plant evolution. Plant J, 72, 947-959.

Doi:10.1111/tpj.12005 pdf

Onnela, J. -P., Fenn, D. J., Reid, S., Porter, M. A., Mucha, P. J., Fricker, M. D.,Jones, N. S. (2012). Taxonomies of networks from community structure. Physical Review E, 86, 036104.

Doi:10.1103/PhysRevE.86.036104 pdf

Schwarzländer, M., Murphy, M. P., Duchen, M. R., Logan, D. C., Fricker, M. D., Halestrap, A. P., Sweetlove, L. J. (2012). Mitochondrial 'flashes': a radical concept repHined. Trends Cell Biol, 22, 503-508.

Doi:10.1016/j.tcb.2012.07.007 pdf

Heaton, L. L., López, E., Maini, P. K., Fricker, M. D., & Jones, N. S. (2012). Advection, diffusion, and delivery over a network. Phys Rev E, 86, 021905.

Doi: http://dx.doi.org/10.1103/PhysRevE.86.021905 pdf

Obara, B., Grau, V., & Fricker, M. D. (2012). A bioimage informatics approach to automatically extract complex fungal networks. Bioinformatics, 28, 2374-2381.

Doi:10.1093/bioinformatics/bts364 pdf

Schwarzländer, M., Logan, D. C., Johnston, I. G., Jones, N. S., Meyer, A. J., Fricker, M. D.,Sweetlove, L. J. (2012). Pulsing of membrane potential in individual mitochondria: a stress-induced mechanism to regulate respiratory bioenergetics in Arabidopsis. Plant Cell, 24, 1188-1201.

Doi:10.1105/tpc.112.096438 pdf

Obara, B., Fricker, M., Gavaghan, D., & Grau, V. (2012). Contrast-independent curvilinear structure detection in biomedical images. IEEE Trans Image Process, 21, 2572-2581.

Doi:10.1109/TIP.2012.2185938 pdf

Au, K. K. C., Pérez-Gõmez, J., Neto, H., Neto, H., Müller, C., Meyer, A. J.,Moore, I. (2012). A perturbation in glutathione biosynthesis disrupts endoplasmic reticulum morphology and secretory membrane traffic in Arabidopsis thaliana. Plant Journal, 71, 881-894.

Doi:10.1111/j.1365-313X.2012.05022.x pdf

Heaton, L., Obara, B., Obara, B., Grau, V., Grau, V., Grau, V.,Fricker, M. D. (2012). Analysis of fungal networks. Fungal Biology Reviews, 26, 12-29.

Doi:10.1016/j.fbr.2012.02.001 pdf

Obara, B., Obara, B., Fricker, M., Grau, V., & Grau, V. (2012). Coherence enhancing diffusion filtering based on the Phase Congruency Tensor. Proceedings - International Symposium on Biomedical Imaging, 202-205.

Doi:10.1109/ISBI.2012.6235519 pdf

Obara, B., Obara, B., Fricker, M., Grau, V., Grau, V., & Grau, V. (2012). Contrast independent detection of branching points in network-like structures. Progress in Biomedical Optics and Imaging - Proceedings of SPIE, 8314.

Doi:10.1117/12.910575 pdf

Obara, B., Obara, B., Fricker, M., Grau, V., & Grau, V. (2012). Local phase approaches to extract biomedical networks. Proceedings - International Symposium on Biomedical Imaging, 1796-1799.

Doi:10.1109/ISBI.2012.6235931 pdf



Schwarzländer, M., Logan, D.C., Fricker, M.D., & Sweetlove, L.J. (2011). The circularly permuted yellow fluorescent protein cpYFP that has been used as a superoxide probe is highly responsive to pH but not superoxide in mitochondria: implications for the existence of superoxide 'flashes'. Biochem J, 437, 381-387.

Doi:10.1042/BJ20110883 pdf

Smith, D.M.D., Onnela, J.P., Onnela, J.P., Lee, C.F., Fricker, M.D.,Johnson, N.F. (2011). Network automata: Coupling structure and function in dynamic networks. Advances in Complex Systems, 14, 317-339.




Heaton, L.L., López, E., Maini, P.K., Fricker, M.D., & Jones, N.S. (2010). Growth-induced mass flows in fungal networks. Proc Roy Soc B Biol Sci, 277, 3265-3274.

Doi:10.1098/rspb.2010.0735 pdf

Boddy, L., Wood, J., Redman, E., Hynes, J., & Fricker, M. D. (2010). Fungal network responses to grazing. Fungal Genet Biol, 47, 522-530.

Doi:10.1016/j.fgb.2010.01.006 pdf

Tero, A., Takagi, S., Saigusa, T., Ito, K., Bebber, D. P., Fricker, M. D.,Nakagaki, T. (2010). Rules for biologically inspired adaptive network design. Science, 327, 439-442.

Doi:10.1126/science.1177894 pdf



Hafke, J. B., Furch, A. C., Fricker, M. D., & van Bel, A. J. (2009). Forisome dispersion in Vicia faba is triggered by Ca2+ hotspots created by concerted action of diverse Ca2+ channels in sieve elements. Plant Signal Behav, 4, 968-972.

Doi: http://dx.doi.org/ 10.4161/psb.4.10.9671 pdf

Furch, A. C., van Bel, A. J., Fricker, M. D., Felle, H. H., Fuchs, M., & Hafke, J. B. (2009). Sieve element Ca2+ channels as relay stations between remote stimuli and sieve tube occlusion in Vicia faba. Plant Cell, 21, 2118-2132.

Doi:10.1105/tpc.108.063107 pdf

Marty, L., Siala, W., Schwarzländer, M., Fricker, M. D., Wirtz, M., Sweetlove, L. J.,Hell, R. (2009). The NADPH-dependent thioredoxin system constitutes a functional backup for cytosolic glutathione reductase in Arabidopsis. PNAS, 106, 9109-9114.

Doi:10.1073/pnas.0900206106 pdf

Schwarzländer, M., Fricker, M. D., & Sweetlove, L. J. (2009). Monitoring the in vivo redox state of plant mitochondria: effect of respiratory inhibitors, abiotic stress and assessment of recovery from oxidative challenge. Biochim Biophys Acta, 1787, 468-475.

Doi:10.1016/j.bbabio.2009.01.020 pdf

Lehmann, M., Schwarzländer, M., Obata, T., Sirikantaramas, S., Burow, M., Olsen, C. E.,Laxa, M. (2009). The metabolic response of Arabidopsis roots to oxidative stress is distinct from that of heterotrophic cells in culture and highlights a complex relationship between the levels of transcripts, metabolites, and flux. Mol Plant, 2, 390-406.

Doi:10.1093/mp/ssn080 pdf

Boddy, L., Hynes, J., Bebber, D. P., & Fricker, M. D. (2009). Saprotrophic cord systems: Dispersal mechanisms in space and time. Mycoscience, 50, 9-19.

Doi:10.1007/s10267-008-0450-4 pdf

Fricker, M.D. Boddy, L. Nakagaki, T. Bebber, D.P. (2009) Adaptive biological networks. In Adaptive Networks: Theory, Models and Applications. Eds T. Gross and H. Sayama, Pp 51-70.

Doi: 10.1007/978-3-642-01284-6_4 pdf



Ingle, R. A., Fricker, M. D., & Smith, J. A. (2008). Evidence for nickel/proton antiport activity at the tonoplast of the hyperaccumulator plant Alyssum lesbiacum. Plant Biol (Stuttg), 10, 746-753.

Doi:10.1111/j.1438-8677.2008.00080.x pdf

Rotheray, T. D., Jones, T. H., Fricker, M. D., & Boddy, L. (2008). Grazing alters network architecture during interspecific mycelial interactions. Fungal Ecology, 1, 124-132.

Doi:10.1016/j.funeco.2008.12.001 pdf

Schwarzländer, M., Fricker, M. D., Müller, C., Marty, L., Brach, T., Novak, J.,Meyer, A. J. (2008). Confocal imaging of glutathione redox potential in living plant cells. J Microsc, 231, 299-316.

Doi:10.1111/j.1365-2818.2008.02030.x pdf

Fricker, M. D., Lee, J. A., Bebber, D. P., Tlalka, M., Hynes, J., Darrah, P. R., Boddy, L. (2008). Imaging complex nutrient dynamics in mycelial networks. J Microsc, 231, 317-331.

Doi:10.1111/j.1365-2818.2008.02043.x pdf

Tlalka, M., Bebber, D. P., Darrah, P. R., Watkinson, S. C., & Fricker, M. D. (2008). Quantifying dynamic resource allocation illuminates foraging strategy in Phanerochaete velutina. Fungal Genet Biol, 45, 1111-1121.

Doi:10.1016/j.fgb.2008.03.015 pdf

Morgan, M. J., Lehmann, M., Schwarzländer, M., Baxter, C. J., Sienkiewicz-Porzucek, A., Williams, T. C.,Finkemeier, I. (2008). Decrease in manganese superoxide dismutase leads to reduced root growth and affects tricarboxylic acid cycle flux and mitochondrial redox homeostasis. Plant Physiol, 147, 101-114.

Doi:10.1104/pp.107.113613 pdf

Tlalka, M., Fricker, M., & Watkinson, S. (2008). Imaging of long-distance alpha-aminoisobutyric acid translocation dynamics during resource capture by Serpula lacrymans. Appl Environ Microbiol, 74, 2700-2708.

Doi:10.1128/AEM.02765-07 pdf

Samalova, M., Fricker, M., & Moore, I. (2008). Quantitative and qualitative analysis of plant membrane traffic using fluorescent proteins. Methods Cell Biol, 85, 353-380.

Doi:10.1016/S0091-679X(08)85015-7 pdf

Meyer, A.J. and Fricker, M.D. (2008). Imaging thiol-based redox processes in live cells. In: Sulfur Metabolism in Phototropic Organisms. Ed: C. Dahl, R. Hell, D. Knaff, T. Leustek. pp. 483–501.

ISBN: 1402068638, 9781402068638 pdf

Fricker, M.D., Bebber, D.P. and Boddy, L. (2008) Mycelial networks: structure and dynamics. In: Ecology of saprotrophic basiodiomycetes. Eds L. Boddy, J.C. Franklin and P. van West. Pp 3-18.

Doi: http://dx.doi.org/10.1016/S0275-0287(08)80003-3 pdf



Tlalka, M., Bebber, D. P., Darrah, P. R., Watkinson, S. C., & Fricker, M. D. (2007). Emergence of self-organised oscillatory domains in fungal mycelia. Fungal Genet Biol, 44, 1085-1095.

Doi:10.1016/j.fgb.2007.02.013 pdf

Fricker, M. D., Tlalka, M., Bebber, D., Takagi, S., Watkinson, S. C., & Darrah, P. R. (2007). Fourier-based spatial mapping of oscillatory phenomena in fungi. Fungal Genet Biol, 44, 1077-1084.

Doi:10.1016/j.fgb.2007.02.012 pdf

Bebber, D. P., Hynes, J., Darrah, P. R., Boddy, L., & Fricker, M. D. (2007). Biological solutions to transport network design. Proc Roy Soc Biol Sci, 274, 2307-2315.

Doi:10.1098/rspb.2007.0459 pdf

Fricker, M.D., Boddy, L. and Bebber, D.P. (2007) Network organisation of mycelial fungi. In: The Mycota. Vol VIII, Biology of the Fungal Cell (2nd Ed). Eds R.J. Howard and N.A.R. Gow. 2nd ed., 341

Doi: http://dx.doi.org/10.1007/978-3-540-70618-2_13 pdf



Libourel, I. G., van Bodegom, P. M., Fricker, M. D., & Ratcliffe, R. G. (2006). Nitrite reduces cytoplasmic acidosis under anoxia. Plant Physiol, 142, 1710-1717.

Doi:10.1104/pp.106.088898 pdf

Samalova, M., Fricker, M., & Moore, I. (2006). Ratiometric fluorescence-imaging assays of plant membrane traffic using polyproteins. Traffic, 7, 1701-1723.

Doi:10.1111/j.1600-0854.2006.00502.x pdf

Bebber, D.P., Tlalka, M., Hynes, J., Darrah, P.R., Ashford, A., Watkinson, S.C., Boddy, L., and Fricker, M.D. (2006). Imaging complex nutrient dynamics in mycelial networks. In: G.M. Gadd (Ed). Fungi in the Environment. Cambridge University Press, Cambridge.

ISBN: 1139462105, 9781139462105 pdf

Watkinson, S.C., Bebber, D., Darrah, P.R., Fricker, M.D.,Tlalka, M. & Boddy, L. (2006) The role of wood decay fungi in the carbon and nitrogen dynamics of the forest floor. In: Gadd, G.M. (ed.) Fungi in Biogeochemical Cycles. Cambridge University Press, Cambridge.

ISBN 1107320747, 9781107320741 pdf

Jarrett, T. C., Ashton, D. J., Fricker, M., & Johnson, N. F. (2006). Interplay between function and structure in complex networks. Phys Rev E, 74, 026116.

Doi: http://dx.doi.org/ http://dx.doi.org/10.1103/PhysRevE.74.026116 pdf

Darrah, P. R., Tlalka, M., Ashford, A., Watkinson, S. C., & Fricker, M. D. (2006). The vacuole system is a significant intracellular pathway for longitudinal solute transport in basidiomycete fungi. Eukaryot Cell, 5, 1111-1125.

Doi:10.1128/EC.00026-06 pdf

Fricker, M., Runions, J., & Moore, I. (2006). Quantitative fluorescence microscopy: from art to science. Annu Rev Plant Biol, 57, 79-107.

Doi:10.1146/annurev.arplant.57.032905.105239 pdf

Boddy, L., Tordoff, G.M., Wood, J., Hynes, J., Bebber, D., Jones, T.H. and Fricker, M.D. (2006). Mycelial foraging strategies of saprotrophic cord-forming basidiomycetes. In: 8th International Mycological Congress Proceedings, ed. W. Meyer & C. Pearce. Medimond, Italy. Pp. 13-20.  pdf



Watkinson, S. C., Boddy, L., Burton, K., Darrah, P. R., Eastwood, D., Fricker, M. D.,Tlalka, M. (2005). New approaches to investigating the function of mycelial networks. Mycologist, 19, 11-17.

Doi:10.1017/S0269915XO5001023 pdf



Meskauskas, A., Fricker, M. D., & Moore, D. (2004). Simulating colonial growth of fungi with the Neighbour-Sensing model of hyphal growth. Mycol Res, 108, 1241-1256.

Doi:http://dx.doi.org/10.1017/S0953756204001261 pdf



Hartmann, T. N., Fricker, M. D., Rennenberg, H., & Meyer, A. J. (2003). Cell-specific measurement of cytosolic glutathione in poplar leaves. Plant Cell Environ, 26, 965-975.

Doi: http://dx.doi.org/ 10.1046/j.1365-3040.2003.01031.x pdf

Tlalka, M., Hensman, D., Darrah, P. R., Watkinson, S. C., & Fricker, M. D. (2003). Noncircadian oscillations in amino acid transport have complementary profiles in assimilatory and foraging hyphae of Phanerochaete velutina. New Phytologist, 158, 325-335.

Doi:10.1046/j.1469-8137.2003.00737.x pdf



Meyer, A. J., & Fricker, M. D. (2002). Control of demand-driven biosynthesis of glutathione in green Arabidopsis suspension culture cells. Plant Physiol, 130, 1927-1937. Doi:10.1104/pp.008243 pdf

Brandizzi, F., Fricker, M., & Hawes, C. (2002). A greener world: the revolution in plant bioimaging. Nat Rev Mol Cell Biol, 3, 520-530.

Doi: 10.1038/nrm861 pdf

Tlalka, M., Watkinson, S. C., Darrah, P. R., & Fricker, M. D. (2002). Continuous imaging of amino-acid translocation in intact mycelia of Phanerochaete velutina reveals rapid, pulsatile fluxes. New Phytologist, 153, 173-184.

Doi:10.1046/j.0028-646X.2001.00288.x pdf



Meyer, A. J., May, M. J., & Fricker, M. (2001). Quantitative in vivo measurement of glutathione in Arabidopsis cells. Plant J, 27, 67-78.

Doi: http://dx.doi.org/10.1046/j.1365-313X.2001.01071.x pdf

Fricker, M. D., & Meyer, A. J. (2001). Confocal imaging of metabolism in vivo: pitfalls and possibilities. J Exp Bot, 52, 631-640.

Doi: http://dx.doi.org/10.1093/jexbot/52.356.631 pdf

Fricker, M.D., Parsons, A., Tlalka, M., Blancaflor, E., Gilroy, S., Meyer, A. and Plieth, C. (2001) Fluorescent probes for living plant cells. In: Plant Cell Biology: A Practical Approach. 2nd Ed. Ed C. Hawes and B. Satiat-Jeunemaitre. Pp 35-84. pdf



Gutiérrez-Alcalá, G., Gotor, C., Meyer, A. J., Fricker, M., Vega, J. M., & Romero, L. C. (2000). Glutathione biosynthesis in Arabidopsis trichome cells. PNAS, 97, 11108-11113.

Doi: http://dx.doi.org/ 10.1073/pnas.190334497 pdf

Meyer, A. J., & Fricker, M. D. (2000). Direct measurement of glutathione in epidermal cells of intact Arabidopsis roots by two-photon laser scanning microscopy. J Microsc, 198, 174-181.

Doi: doi:10.1046/j.1365-2818.2000.00696.x" pdf

Fricker, M. D., May, M., Meyer, A. J., Sheard, N., & White, N. S. (2000). Measurement of glutathione levels in intact roots of Arabidopsis. J Microsc, 198, 162-173.

Doi: 10.1046/j.1365-2818.2000.00696.x pdf

Fricker, M.D., May, M., White, N.S. and Meyer, A. (2000) Modelling GSH pathways from in vivo imaging. In: Sulfur Nutrition and Sulfur Assimilation in Higher Plants. Ed. C. Brunold, H. Rennenberg, L.J. De Kok, I. Stulen and J.-C. Davidan. Paul Haupt Publishers, Berne. Pp 353-354.

Meyer, A.J. and Fricker, M.D. (2000) Quantitative imaging of glutathione in live cells by laser scanning microscopy. In: Sulfur Nutrition and Sulfur Assimilation in Higher Plants. Ed. C. Brunold, H. Rennenberg, L.J. De Kok, I. Stulen and J.-C. Davidan. Paul Haupt Publishers, Berne. Pp 351-352.



Tlałka, M., Runquist, M., Runquist, M., & Fricker, M. (1999). Light perception and the role of the xanthophyll cycle in blue-light-dependent chloroplast movements in Lemna trisulca L. Plant Journal, 20, 447-459.

Doi:10.1046/j.1365-313X.1999.00614.x pdf

Tlałka, M., & Fricker, M. (1999). The role of calcium in blue-light-dependent chloroplast movement in Lemna trisulca L. Plant Journal, 20, 461-473.

Doi:10.1046/j.1365-313X.1999.00621.x pdf

Fricker, M.D. and Oparka, K.J. (1999) Imaging techniques in plant transport: Meeting review. J. Exp. Bot. 50 1089-1100.

URL: http://www.jstor.org/stable/23696211 pdf

Fricker, M.D., Plieth, C., Knight, H., Blancaflor, E., Knight, M.R., White, N.S., and Gilroy, S. (1999) Fluorescent and luminescent techniques to probe ion activities in living plant cells. In: Fluorescent and luminescent probes. 2nd Ed. Ed. W.T. Mason. Pp 569-596.

Doi: 10.1016/B978-012447836-7/50044-0 pdf



Doe, C. L., Wang, G., Chow, C., Fricker, M. D., Singh, P. B., & Mellor, E. J. (1998). The fission yeast chromo domain encoding gene chp1+ is required for chromosome segregation and shows a genetic interaction with alpha-tubulin. Nucleic Acids Res, 26, 4222-4229.

Doi: http://dx.doi.org/10.1093/nar/26.18.4222 pdf



Fricker, M. D., White, N. S., & Obermeyer, G. (1997). pH gradients are not associated with tip growth in pollen tubes of Lilium longiflorum. J Cell Sci, 110, 1729-1740.

URL: http://www.ncbi.nlm.nih.gov/pubmed/9264460 pdf

Sánchez-Fernández, R., Fricker, M., Corben, L. B., White, N. S., Sheard, N., Leaver, C. J.,May, M. J. (1997). Cell proliferation and hair tip growth in the Arabidopsis root are under mechanistically different forms of redox control. PNAS, 94, 2745-2750.

Doi:10.1073/pnas.94.6.2745 pdf

Errington, R. J., Fricker, M. D., Wood, J. L., Hall, A. C., & White, N. S. (1997). Four-dimensional imaging of living chondrocytes in cartilage using confocal microscopy: a pragmatic approach. Am J Physiol, 272, C1040-C1051.

URL: http://ajpcell.physiology.org/content/272/3/C1040.short pdf

Fricker, M., Hollinshead, M., White, N., & Vaux, D. (1997). Interphase nuclei of many mammalian cell types contain deep, dynamic, tubular membrane-bound invaginations of the nuclear envelope. J Cell Biol, 136, 531-544.

Doi: http://dx.doi.org/ 10.1083/jcb.136.3.531 pdf

Fricker, M.D., Errington, R.J., Wood, J.L., Tlalka, M., May, M. and White, N.S. (1997) Quantitative confocal fluorescence measurements in living tissues. In: Signal Transduction - Single Cell Research. Ed. B. Van Duijn. and A. Wiltnik. Springer-Verlag, Heidelberg. 413-445.

Doi: http://dx.doi.org/10.1007/s00898-997-0019-2 pdf

White, N.S., Errington, R.J., Fricker, M.D. and Wood, J.L. (1997) Multidimensional fluorescence microscopy: optical distortions in quantitative imaging of biological specimens. In: Fluorescence Microscopy and Fluorescent Probes. Plenum Press.

Doi: http://dx.doi.org/10.1007/978-1-4899-1866-6_4



Willmer, CM, Fricker, MD. (1996) Stomata. 2nd Ed. 

ISBN: 0412574306, 9780412574306

White, N. S., Errington, R. J., Fricker, M. D., & Wood, J. L. (1996). Aberration control in quantitative imaging of botanical specimens by multidimensional fluorescence microscopy. Journal of Microscopy, 181, 99-116.

Doi:10.1046/j.1365-2818.1996.113392.x pdf



Napier, R., Trueman, S., Henderson, J., Boyce, J., Hawes, C., Fricker, M.,Venis, M. (1995). Purification, sequencing and functions of calreticulin from maize. J. Exp.Botany, 46, 1603-1613.

Doi:10.1093/jxb/46.10.1603 pdf



Boyce, J. M., Coates, D., Fricker, M. D., & Evans, D. E. (1994). Genomic sequence of a calnexin homolog from Arabidopsis thaliana. Plant Physiol, 106, 1691.

URL: http://www.ncbi.nlm.nih.gov/pmc/articles/PMC159718/ pdf

Larin, Z., Fricker, M. D., Maher, E., Ishikawa-Brush, Y., & Southern, E. M. (1994). Fluorescence in situ hybridisation of multiple probes on a single microscope slide. Nucleic Acids Res, 22, 3689-3692.

Doi: http://dx.doi.org/10.1093/nar/22.18.3689 pdf

Allan, A. C., Fricker, M. D., Ward, J. L., Beale, M. H., & Trewavas, A. J. (1994). Two transduction pathways mediate rapid effects of abscisic acid in Commelina guard cells. Plant Cell, 6, 1319-1328.

Doi:10.1105/tpc.6.9.1319 pdf

Larin, Z., Fricker, M. D., & Tyler-Smith, C. (1994). De novo formation of several features of a centromere following introduction of a Y alphoid YAC into mammalian cells. Hum Mol Genet, 3, 689-695.

Doi: http://www.ncbi.nlm.nih.gov/pubmed/8081354 pdf

Fricker, M.D., White, N.S., Thiel, G., Millner, P., and Blatt, M.R. (1994) Peptides derived from the auxin binding protein elevate Ca2+ and pH in stomatal guard cells of Vicia: A confocal fluorescence ratio imaging study. In: Membrane Transport in Plants and fungi: Molecular Mechanisms and Contol. Eds. M.R. Blatt, R.A. Leigh and D. Sanders. SEB Symposium Series 48 215-228.

Fricker, MD, Tlalka, M, Ermantraut, J, Obermeyer, G, Dewey, M, Gurr, S, Patrick, J, White, NS. (1994) Confocal fluorescence ratio imaging of ion activities in plant cells Scanning Microscopy. 8: pp 391-405. 



Thiel, G., Blatt, M. R., Fricker, M. D., White, I. R., & Millner, P. (1993). Modulation of K+ channels in Vicia stomatal guard cells by peptide homologs to the auxin-binding protein C terminus. PNAS, 90, 11493-11497.

Doi: doi:10.1073/pnas.90.24.11493 pdf

Fricker, M.D. Blatt, M.R. and White, N.S. (1993) Confocal fluorescence ratio imaging of pH in plant cells. In: Biotechnology Applications of Microinjection, Microscopic Imaging and Fluorescence. Eds. P. Bach, C.H. Reynolds, J.M. Clark, P.L. Poole and J. Mottley Plenum Press. Pp 151-162.

Doi: http://dx.doi.org/10.1007/978-1-4615-2828-9_18

Fricker, M.D., Tester, M. and Gilroy, S. (1993) Fluorescent and luminescent techniques to probe plant cell activity. In W.T. Mason and G. Relf, Eds, Fluorescent Probes for Biological Activity of Living Cells - A Practical guide. Academic Press. Pp 360-377.



Fricker, M. D., & White, N. S. (1992). Wavelength considerations in confocal microscopy of botanical specimens. J. Microscopy, 166(1), 29-42.

Doi: http://dx.doi.org/ 10.1111/j.1365-2818.1992.tb01505.x pdf

Fricker, M.D. and White, N.S. (1992) Application of confocal microscopy and three-dimensional image analysis to plant and microbial cells. Binary 4 44-49.



Hodick, D., Gilroy, S., Fricker, M., & Trewavas, A. (1991). Cytosolic Ca2+-concentrations and distributions in rhizoids of Chara fragilis desv. determined by ratio analysis of the fluorescent-probe indo-1. Botanica Acta, 104, 222-228.

Doi: http://dx.doi.org/ 10.1111/j.1438-8677.1991.tb00221.x

Fricker, M. D., Grantz, D. A., & Willmer, C. M. (1991). Stomatal responses measured using a viscous flow (liquid) porometer. J. Exp. Bot., 42, 747-755.

Doi:10.1093/jxb/42.6.747 pdf

Gilroy, S., Fricker, M. D., Read, N. D., & Trewavas, A. J. (1991). Role of calcium in signal transduction of Commelina guard cells. Plant Cell, 3, 333-344.

Doi:10.1105/tpc.3.4.333 pdf

Fricker, M.D., Gilroy, S.G., Read, N.D. and Trewavas, A.J. (1991) Visualisation and measurement of the calcium message in guard cells. In: Molecular Biology of Plant Development. Eds: G.I. Jenkins and W. Schuch. SEB Symposium Series, 44, 177-190.

Gilroy, S., Fricker, M.D., Read, N.D. and Trewavas, A.J. (1991) The role of Ca2+ and ABA in the regulation of stomatal aperture. In: Progress in Plant Growth Regulation. Eds. C.M. Karssen, L.C. Van Loon and D. Vreugdenhil. Current Plant Science and Biotechnology in Agriculture, 13, 105-115. Kluwer Academic Publishers. Dordrecht. 

White, N.S., Bennett, S.T., Kenton, A.Y., Callimassia, M.A. and Fricker, M.D. (1991) Characterising plant chromosomes and their 3-D organisation using CLSM. Scanning 13, 228-229.

White, N.S., Fricker, M.D. and Shotton, D.M. (1991) Quantitative visualisation of 3D biological CLSM images. Scanning 13, 51-53.



Fricker, M., & Willmer, C. (1990). Nitrate-sensitive ATPase activity and proton pumping in guard cell protoplasts of Commelina. J. Exp. Bot, 41, 193-198.

Doi:10.1093/jxb/41.2.193 pdf

Bennett, S.T., Fricker, M.D., Bennett, M.D. and White, N.S. (1990) The 3-D localization of chromosomes using confocal microscopy. Trans. Royal Microscopical Soc. 1, 441-444.

Fricker, M.D., Gilroy, S.G. and Trewavas, A.J. (1990). Signal transduction in plant cells and the calcium message. In: Signal Perception and Transduction in Higher Plants. NATO AS1 Series, 47, pp 89-102. Eds R. Ranjeva and A.M. Boudet. Springer-Verlag Berlin Heidelberg.

Fricker, M.D. and White, N.S. (1990) Volume measurement of guard cell vacuoles during stomatal movements using confocal microscopy. Trans. Royal Microscopical Soc. 1, 345-348. pdf

Gilroy, S., Fricker, M.D., Read, N.D. and Trewavas, A.J. (1990) Fluorescence ratio imaging and photometry of calcium in living plant cells. Trans. Royal Microscopical Soc. 1, 475-478.



Gilroy, S.G., Fricker, M.D., Blowers, D., Harvey, H.J., Collinge, M. and Trewavas, A.J. (1989) Calcium channels, cytosol calcium and plasma membrane phosphorylation: an integrated calcium stat system. In: Plant Membrane Transport: The Current Position. Pp 215-224. Eds J. Dainty, M.I. DeMichelis, E. Marre and F. Rasi-Caldogno. Elsevier. Amsterdam.



Fricker, M., & Willmer, C. (1987). Vanadate sensitive ATPase and phosphatase-activity in guard-cell protoplasts of Commelina. J. Exp. Bot, 38, 642-648.

Doi:10.1093/jxb/38.4.642 pdf