Institute of Neuroscience and Psychology

Prof Andrew Todd

  • Professor of Neuroscience (Neurosciences and Clinical Psychology)

telephone: 01413305868
email: Andrew.Todd@glasgow.ac.uk


Neurochemistry and Synaptic Connections in the Mammalian Spinal Cord
My main research interest is the organisation of neuronal circuits in the spinal dorsal horn that underlie the perception of pain and itch. We use immunocytochemistry and confocal microscopy combined with other techniques, such as neuronal tract-tracing, patch-clamp recording from spinal cord slices, and electron microscopy. I have collaborations with: Dr Marco Beato (UCL); Dr Toshi Yasaka (Saga University, Japan), Dr Sarah Ross (University of Pittsburgh, USA); Dr Peter Szucs and Prof Boris Safronov (University of Porto, Portugal); Dr Masahiko Watanabe (Hokkaido University, Japan). My work is funded by the Wellcome Trust and the BBSRC.

Organisation of neuronal circuits in the dorsal horn
Although the dorsal horn plays a major role in transmitting and modifying incoming sensory information, we know little about the nerve circuits responsible for these functions, largely due to the difficulty in defining functional populations of neurons.

Projection neurons
Projection neurons, with axons that travel directly to the brain, are concentrated in lamina I, and most of these express the neurokinin 1 receptor (NK1r). There are ~400 lamina I projection neurons per side in the rat L4 segment, corresponding to ~5% of all neurons in this lamina. Many project to more than one brain region. There are also  NK1r-expressing projection neurons in lamina III, with similar supraspinal targets, and we have identified a population of giant lamina I projection cells that lack the NK1r.

 

Interneurons
Most neurons in laminae I-III have axons that remain in the spinal cord and are defined as interneurons. They can be divided into two main groups: excitatory and inhibitory. We have shown that among the inhibitory interneurons 4 populations can be defined by the expression of neuropeptide Y (NPY), galanin/dynorphin, neuronal nitric oxide synthase (nNOS) and parvalbumin. These account for over half the inhibitory interneurons in laminae I-II.

 

We have shown that they differ in their postsynaptic targets, e.g. NPY/GABA axons preferentially innervate NK1r+ lamina III projection cells, while nNOS/GABA axons innervate lamina I giant cells. Some of the neuronal circuits that have been identified in laminae I-III are summarised below.

 

Changes in dorsal horn following nerve injury
Injury to peripheral nerves often leads to neuropathic pain, but the underlying mechanisms are poorly understood. There is evidence that changes in the dorsal horn contribute to certain types of neuropathic pain, and several possible mechanisms have been proposed. These include: (1) sprouting of Aβ low-threshold mechanoreceptive afferents into the superficial laminae, where they would have access to nociceptive-specific projection neurons; (2) a phenotypic switch, such that these afferents start to synthesise substance P, which could activate NK1rs on nociceptive projection neurons; (3) loss of inhibitory interneurons from the superficial laminae.

We have provided evidence against each of these suggestions. We found that both before and after nerve injury, the central projections of Aβ afferents extend into, but not beyond the inner half of lamina II. We also showed that the central terminals of these afferents did not express or release substance P after nerve injury. We found that in rat models of neuropathic pain, there was no change in the total number of neurons in laminae I-III, nor in the proportion that were GABA-immunoreactive. There is known to be apoptotic cell death in the dorsal horn in these models, as shown by staining with the TUNEL (Terminal deoxynucleotidyl transferase dUTP nick end labeling) method. However, we showed that this was restricted to microglia (labelled with Iba1 antibody), and that TUNEL positive cells were not neurons.

 

Interneurons in the ventral horn
Although most of our work is on the dorsal horn, we have also identified two important populations of ventral horn interneurons: those giving rise to the "P" and "C" boutons that are associated with motoneurons.

P boutons are GABAergic terminals that form axoaxonic synapses on Ia muscle spindle afferents and regulate proprioceptive input to motoneurons. We showed that P boutons were unique in expressing the 65kDa molecular weight form of glutamic acid decarboxylase (GAD65) This enabled us to identify their cells of origin, which were in the deep medial dorsal horn, and showed that P boutons are likely to represent the only output for these cells.

The C boutons are cholinergic axons that regulate motoneuronexcitability through m2 muscarinic receptors. In collaboration with Drs Rob Brownstone and Gareth Miles (Dalhousie University, Canada) we used a neurochemical/developmental approach to demonstrate that these boutons originate from a group of cholinergic cells located lateral to the central canal (cc) – the so-called "medial partition cells". This was based on the finding that the C boutons, which express choline acetyltransferase (ChAT), lack neuronal nitric oxide synthase (nNOS) but transiently express the transcription factor Dbx1, which was captured with yellow fluorescent protein (YFP).

 

Jump to: 2012 | 2011 | 2010 | 2009 | 2008 | 2007 | 2006 | 2005 | 2004 | 2003 | 2002 | 2001
Number of items: 58.

2012

Baseer, N., Watanabe, M., Furuta, T., Kaneko, T., and Todd, A.J. (2012) Projection neurons in lamina III of the rat spinal cord are selectively innervated by local dynorphin-containing excitatory neurons. Journal of Neuroscience, 32 (34). pp. 11854-11863. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.2707-12.2012)

Tappe-Theodor, A. et al. (2012) Gαq/11 signaling tonically modulates nociceptor function and contributes to activity-dependent sensitization. Pain, 153 (1). pp. 184-196. ISSN 0304-3959 (doi:10.1016/j.pain.2011.10.014)

Baseer, N., Polgar, E., Watanabe, M., Furuta, T., Kaneko, T., and Todd, A. (2012) Projection neurons in Lamina III of the rat spinal cord are selectively innervated by local dynorphin-containing excitatory neurons. Journal of Neuroscience, 32 (34). pp. 11854-11863. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.2707-12.2012)

Luo, C. et al. (2012) Presynaptically localized cyclic GMP-dependent protein kinase 1 is a key determinant of spinal synaptic potentiation and pain hypersensitivity. PLoS Biology, 10 (3). e1001283. ISSN 1544-9173 (doi:10.1371/journal.pbio.1001283)

Todd, A.J. (2012) How to recognise collateral damage in partial nerve injury models of neuropathic pain. Pain, 153 (1). pp. 11-12. ISSN 0304-3959 (doi:10.1016/j.pain.2011.10.031)

2011

Polgár, E., Sardella, T.C.P., Watanabe, M., and Todd, A.J. (2011) Quantitative study of NPY-expressing GABAergic neurons and axons in rat spinal dorsal horn. Journal of Comparative Neurology, 519 (6). pp. 1007-1023. ISSN 0021-9967 (doi:10.1002/cne.22570)

Saywell, S.A., Ford, T.W., Meehan, C.F., Todd, A.J., and Kirkwood, P.A. (2011) Electrophysiological and morphological characterization of propriospinal interneurons in the thoracic spinal cord. Journal of Neurophysiology, 105 (2). pp. 806-826. ISSN 0022-3077 (doi:10.1152/jn.00738.2010)

Gangadharan, V. et al. (2011) Peripheral calcium-permeable AMPA receptors regulate chronic inflammatory pain in mice. Journal of Clinical Investigation, 121 (4). pp. 1608-1623. ISSN 0021-9738 (doi:10.1172/JCI44911)

Sardella, T.C.P., Polgar, E., Watanabe, M., and Todd, A.J. (2011) A quantitative study of neuronal nitric oxide synthase expression in laminae I–III of the rat spinal dorsal horn. Neuroscience, 192 . pp. 708-720. ISSN 0306-4522 (doi:10.1016/j.neuroscience.2011.07.011)

Sardella, T.C.P., Polgár, E., Garzillo, F., Furuta, T., Kaneko, T., Watanabe, M., and Todd, A.J. (2011) Dynorphin is expressed primarily by GABAergic neurons that contain galanin in the rat dorsal horn. Molecular Pain, 7 (1). p. 76. ISSN 1744-8069 (doi:10.1186/1744-8069-7-76)

Tiong, S.Y.X., Polgar, E., van Kralingen, J.C., Watanabe, M., and Todd, A.J. (2011) Galanin-immunoreactivity identifies a distinct population of inhibitory interneurons in laminae I-III of the rat spinal cord. Molecular Pain, 7 (36). pp. 1-13. ISSN 1744-8069 (doi:10.1186/1744-8069-7-36)

2010

Todd, A.J. (2010) Neuronal circuitry for pain processing in the dorsal horn. Nature Reviews Neuroscience, 11 (12). pp. 823-836. ISSN 1471-003X (doi:10.1038/nrn2947)

Polgár, E., Al Ghamdi, K.S., and Todd, A.J. (2010) Two populations of neurokinin 1 receptor-expressing projection neurons in lamina I of the rat spinal cord that differ in AMPA receptor subunit composition and density of excitatory synaptic input. Neuroscience, 167 (4). pp. 1192-1204. ISSN 0306-4522 (doi:10.1016/j.neuroscience.2010.03.028)

Polgár, E., Wright, L.L., and Todd, A.J. (2010) A quantitative study of brainstem projections from lamina I neurons in the cervical and lumbar enlargement of the rat. Brain Research, 1308 . pp. 58-67. ISSN 0006-8993 (doi:10.1016/j.brainres.2009.10.041)

Yasaka, T., Tiong, S.Y.X., Hughes, D.I., Riddell, J.S., and Todd, A.J. (2010) Populations of inhibitory and excitatory interneurons in lamina II of the adult rat spinal dorsal horn revealed by a combined electrophysiological and anatomical approach. Pain, 151 (2). pp. 475-488. ISSN 0304-3959 (doi:10.1016/j.pain.2010.08.008)

2009

Yasaka, T., Hughes, D.I., Polgár, E., Nagy, G.G., Watanabe, M., Riddell, J.S., and Todd, A.J. (2009) Evidence against AMPA receptor-lacking glutamatergic synapses in the superficial dorsal horn of the rat spinal cord. Journal of Neuroscience, 29 (42). pp. 13401-13409. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.2628-09.2009)

Al Ghamdi, K.S., Polgár, E., and Todd, A. (2009) Soma size distinguishes projection neurons from neurokinin 1 receptor-expressing interneurons in lamina I of the rat lumbar spinal dorsal horn. Neuroscience, 164 (4). pp. 1794-1804. ISSN 0306-4522 (doi:10.1016/j.neuroscience.2009.09.071)

Al-Khater, K.M., and Todd, A.J. (2009) Collateral projections of neurons in laminae I, III, and IV of rat spinal cord to thalamus, periaqueductal gray matter, and lateral parabrachial area. Journal of Comparative Neurology, 515 (6). pp. 629-646. ISSN 0021-9967 (doi:10.1002/cne.22081)

Todd, A.J., Polgár, E., Watt, C., Bailey, M.E.S., and Watanabe, M. (2009) Neurokinin 1 receptor-expressing projection neurons in laminae III and IV of the rat spinal cord have synaptic AMPA receptors that contain GluR2, GluR3 and GluR4 subunits. European Journal of Neuroscience, 29 (4). pp. 718-726. ISSN 0953-816X (doi:10.1111/j.1460-9568.2009.06633.x)

2008

Polgár, E., and Todd, A. (2008) Tactile allodynia can occur in the spared nerve injury model in the rat without selective loss of GABA or GABAA receptors from synapses in laminae I–II of the ipsilateral spinal dorsal horn. Neuroscience, 156 (1). pp. 193-202. ISSN 0306-4522 (doi:10.1016/j.neuroscience.2008.07.009)

Polgar, E., Watanabe, M., Hartmann, B., Grant, S., and Todd, A.J. (2008) Expression of AMPA receptor subunits at synapses in laminae I-III of the rodent spinal dorsal horn. Molecular Pain, 4 (5). pp. 1-14. ISSN 1744-8069 (doi:10.1186/1744-8069-4-5)

Al-Khater, K.M., Kerr, R., and Todd, A.J. (2008) A quantitative study of spinothalamic neurons in laminae I, III, and IV in lumbar and cervical segments of the rat spinal cord. Journal of Comparative Neurology, 511 (1). pp. 1-18. ISSN 0021-9967 (doi:10.1002/cne.21811)

Polgar, E., Al-Khater, K. M., Shehab, S., Watanabe, M., and Todd, A.J. (2008) Large Projection Neurons in Lamina I of the Rat Spinal Cord That Lack the Neurokinin 1 Receptor Are Densely Innervated by VGLUT2-Containing Axons and Possess GluR4-Containing AMPA Receptors. Journal of Neuroscience, 28 (49). pp. 13150-13160. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.4053-08.2008)

2007

Hughes, D.I., Scott, D.T., Riddell, J.S., and Todd, A.J. (2007) Up-regulation of substance P in low-threshold myelinated afferents is not required for tactile allodynia in the chronic constriction injury and spinal nerve ligation models. Journal of Neuroscience, 27 (8). pp. 2035-2044. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.5401-06.2007)

Marvizón, J.C.G., Pérez, O.A., Song, B., Chen, W., Bunnett, N.W., Grady, E.F., and Todd, A.J. (2007) Calcitonin receptor-like receptor and receptor activity modifying protein 1 in the rat dorsal horn: localization in glutamatergic presynaptic terminals containing opioids and adrenergic α2C receptors. Journal of Neuroscience, 148 (1). pp. 250-265. ISSN 0270-6474 (doi:10.1016/j.neuroscience.2007.05.036)

Miles, G.B., Hartley, R., Todd, A.J., and Brownstone, R.M. (2007) Spinal cholinergic interneurons regulate the excitability of motoneurons during locomotion. Proceedings of the National Academy of Sciences of the United States of America, 104 (7). pp. 2448-2453. ISSN 1091-6490

Polgár, E., Campbell, A.D., MacIntyre, L., Watanabe, M., and Todd, A.J. (2007) Phosphorylation of ERK in neurokinin 1 receptor-expressing neurons in laminae III and IV of the rat spinal dorsal horn following noxious stimulation. Molecular Pain, 3 (4). ISSN 1744-8069

Hughes, D.I., Scott, D.T., Riddell, J.S., and Todd, A.J. (2007) Upregulation of substance P in low-threshold myelinated afferents is not required for tactile allodynia in the chronic constriction injury and spinal nerve ligation models. Journal of Neuroscience, 27 (8). pp. 2035-2044. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.5401-06.2007)

Marvizon, JCG, Perez, OA, Song, B, Chen, W, Bunnett, NW, Grady, EF, and Todd, AJ (2007) Calcitonin receptor-like and receptor activity modifying protein 1 in the rat dorsal horn localization in glutamatergic presynaptic terminal containing opioids and adrenergic alpha(2c) receptor. Neuroscience, 148 . pp. 250-265. (doi:10.1016/j.neuroscience.2007.05.036)

Polgar, E, Thomson, S, Maxwell, DJ, Al-Khater, K, and Todd, AJ (2007) A population of large neurons in laminae III and IV of the rat spinal cord that have long dorsal dendrites and lack the neurokinin 1 receptor. European Journal of Neuroscience, 26 . pp. 1587-1598. (doi:10.1111/j.1460-9568.2007.05793.x)

2006

Polgar, E, Furuta, T, Kaneko, T, and Todd, A (2006) Characterization of neurons that express preprotachykinin B in the dorsal horn of the rat spinal cord. Neuroscience, 139 . pp. 687-697. (doi:10.1016/j.neuroscience.2005.12.021)

Todd, A (2006) Erratum to 'Characterization of neurons that express preprotachykinin B in the dorsal horn of the rat spinal cord': [Neuroscience 139 (2006) 687-697]. Neuroscience, 141 (4). pp. 2195-2197. (doi:10.1016/j.neuroscience.2005.12.0)

2005

Shehab, S., McGonigle, D., Hughes, D.I., Todd, A.J., and Redgrave, P. (2005) Anatomical evidence for an anticonvulsant relay in the rat ventromedial medulla. European Journal of Neuroscience, 22 (6). pp. 1431-1444. ISSN 0953-816X (doi:10.1111/j.1460-9568.2005.04326.x)

Polgar, E., Hughes, D.I., Arham, A.Z., and Todd, A.J. (2005) Loss of neurons from laminas I-III of the spinal dorsal horn is not required for development of tactile allodynia in the spared nerve injury model of neuropathic pain. Journal of Neuroscience, 25 (28). pp. 6658-6666. ISSN 0270-6474 (doi:10.1523/JNEUROSCI.1490-05.2005)

Hughes, D.I. et al. (2005) P boutons in lamina IX of the rodent spinal cord express high levels of glutamic acid decarboxylase-65 and originate from cells in deep medial dorsal horn. Proceedings of the National Academy of Sciences, 102 (25). pp. 9038-9043. ISSN 0027-8424 (doi:10.1073/pnas.0503646102)

Alvarez, FJ, Jonas, PC, Sapir, T, Hartley, R, Berrocal, MC, Geiman, EJ, Todd, AJ, and Goulding, M (2005) Postnatal phenotype and localization of spinal cord V1 derived interneurons. Journal of Comparative Neurology, 493 . pp. 177-192. (doi:10.1002/cne.20711)

Hinckley, CA, Hartley, R, Wu, L, Todd, A, and Ziskind-Conhaim, L (2005) Locomotor-like rhythms in a genetically distinct cluster of interneurons in the mammalian spinal cord. Journal of Neurophysiology, 93 . pp. 1439-1449. (doi:10.1152/jn.00647.2004)

Todd, AJ, Spike, RC, Young, S, and Puskar, Z (2005) Fos induction in lamina I projection neurons in response to noxious thermal stimuli. Neuroscience, 131 . pp. 209-217. (doi:10.1016/j.neuroscience.2004.11.001)

Wilson, JM, Hartley, R, Maxwell, DJ, Todd, AJ, Lieberam, I, Kaltschmidt, JA, Yoshida, Y, Jessell, TM, and Brownstone, RM (2005) Conditional rhythmicity of ventral spinal interneurons defined by expression of the Hb9 homeodomain protein. Journal of Neuroscience, 25 . pp. 5710-5719. (doi:10.1523/JNEUROSCI.0274-05.2005)

Worsley, MA, Todd, AJ, and King, AE (2005) Serotoninergic-mediated inhibition of substance P sensitive deep dorsal horn neurons: a combined electrophysiological and morphological study in vitro. Experimental Brain Research, 160 . pp. 360-367. (doi:10.1007/s00221-004-2018-7)

2004

Hughes, D.I., Polgár, E., Shehab, S.A.S., and Todd, A.J. (2004) Peripheral axotomy induces depletion of the vesicular glutamate transporter VGLUT1 in central terminals of myelinated afferent fibres in the rat spinal cord. Brain Research, 1017 (1-2). pp. 69-76. ISSN 0006-8993 (doi:10.1016/j.brainres.2004.05.054)

Halstead, SK, OHanlon, GM, Humphreys, PD, Morrison, DB, Morgan, BP, Todd, AJ, Plomp, JJ, and Willison, HJ (2004) Anti-disialoside antibodies kill perisynaptic Schwann cells and damage motor nerve terminals via membrane attack complex in a murine model of neuropathy. Brain, 127 . pp. 2109-2123. (doi:10.1093/brain/awh231)

Nagy, GG, Al-Ayyan, M, Andrew, D, Fukaya, M, Watanabe, M, and Todd, AJ (2004) Widespread expression of the AMPA receptor GluR2 subunit at glutamatergic synapses in the rat spinal cord and phosphorylation of GluR1 in response to noxious stimulation revealed with an antigen-unmasking method. Journal of Neuroscience, 24 . pp. 5766-5777. (doi:10.1523/JNEUROSCI.1237-04.2004)

Nagy, GG, Watanabe, M, Fukaya, M, and Todd, AJ (2004) Synaptic distribution of the NR1, NR2A and NR2B subunits of the N-methyl-D-aspartate receptor in the rat lumbar spinal cord revealed with an antigen-unmasking technique. European Journal of Neuroscience, 20 . pp. 3301-3312. (doi:10.1111/j.1460-9568.2004.03798.x)

Polgar, E, Gray, S, Riddell, JS, and Todd, AJ (2004) Lack of evidence for significant neuronal loss in laminae I-III of the spinal dorsal horn of the rat in the chronic constriction injury model. Pain, 111 . pp. 144-150. (doi:10.1016/j.pain.2004.06.011)

Shehab, SAS, Spike, RC, and Todd, AJ (2004) Do central terminals of intact myelinated primary afferents sprout into the superficial dorsal horn of rat spinal cord after injury to a neighboring peripheral nerve? Journal of Comparative Neurology, 474 . pp. 427-437. (doi:10.1002/cne.20147)

2003

Hughes, D.I., Scott, D.T., Todd, A.J., and Riddell, J.S. (2003) Lack of evidence for sprouting of Aβ afferents into the superficial laminas of the spinal cord dorsal horn after nerve section. Journal of Neuroscience, 23 (29). pp. 9491-9499. ISSN 0270-6474

Polgár, E., Hughes, D.I., Riddell, J.S., Maxwell, D.J., Puskár, Z., and Todd, A.J. (2003) Selective loss of spinal GABAergic or glycinergic neurons is not necessary for development of thermal hyperalgesia in the chronic constriction injury model of neuropathic pain. Pain, 104 (1). pp. 229-239. ISSN 0304-3959 (doi:10.1016/S0304-3959(03)00011-3)

Todd, A.J., Hughes, D.I., Polgar, E., Nagy, G.G., Mackie, M., Ottersen, O.P., and Maxwell, D.J. (2003) The expression of vesicular glutamate transporters VGLUT1 and VGLUT2 in neurochemically defined axonal populations in the rat spinal cord with emphasis on the dorsal horn. European Journal of Neuroscience, 17 (1). pp. 13-27. ISSN 0953-816X (doi:10.1046/j.1460-9568.2003.02406.x)

Hughes, DI, Scott, DT, Todd, AJ, and Riddell, JS (2003) Lack of evidence for sprouting of A beta afferents into the superficial laminas of the spinal cord dorsal horn after nerve section. Journal of Neuroscience, 23 . pp. 9491-9499.

Mackie, M., Hughes, D.I., Maxwell, D.J., Tillakaratne, N.J.K., and Todd, A.J. (2003) Distribution and colocalisation of glutamate decarboxylase isoforms in the rat spinal cord. Neuroscience Letters, 119 . pp. 461-472. ISSN 0304-3940 (doi:10.1016/S0306-4522(03)00174-X)

Shehab, SAS, Spike, RC, and Todd, AJ (2003) Evidence against cholera toxin B subunit as a reliable tracer for sprouting of primary afferents following peripheral nerve injury. Brain Research, 964 . pp. 218-227. (doi:10.1016/S0006-8993(02)04001-5)

Spike, RC, Puskar, Z, Andrew, D, and Todd, AJ (2003) A quantitative and morphological study of projection neurons in lamina I of the rat lumbar spinal cord. European Journal of Neuroscience, 18 . pp. 2433-2448. (doi:10.1046/j.1460-9568.2003.02981.x)

2002

Polgar, E, Puskar, Z, Watt, C, Matesz, C, and Todd, AJ (2002) Selective innervation of lamina I projection neurones that possess the neurokinin 1 receptor by serotonin-containing axons in the rat spinal cord. Neuroscience, 109 . pp. 799-809.

Spike, R.C., Puskar, Z., Sakamoto, H., Stewart, W., Watt, C., and Todd, A.J. (2002) MOR-1-immunoreactive neurons in the dorsal horn of the rat spinal cord: evidence for nonsynaptic innervation by substance P-containing primary afferents and for selective activation by noxious thermal stimuli. European Journal of Neuroscience, 15 (8). pp. 1306-1316. ISSN 0953-816X (doi:10.1046/j.1460-9568.2002.01969.x)

Todd, AJ (2002) Anatomy of primary afferents and projection neurones in the rat spinal dorsal horn with particular emphasis on substance P and the neurokinin 1 receptor. Experimental Physiology, 87 . pp. 245-249.

Todd, AJ, Puskar, Z, Spike, RC, Hughes, C, Watt, C, and Forrest, L (2002) Projection neurons in lamina I of rat spinal cord with the neurokinin 1 receptor are selectively innervated by substance p-containing afferents and respond to noxious stimulation. Journal of Neuroscience, 22 . pp. 4103-4113.

2001

Puskar, Z, Polgar, E, and Todd, AJ (2001) A population of large lamina I projection neurons with selective inhibitory input in rat spinal cord. Neuroscience, 102 . pp. 167-176.

This list was generated on Tue May 21 21:58:12 2013 BST.
  • Wellcome Open Access Award
    Wellcome Trust
    2013 - 2013
     
  • Populations of inhibitory interneurons in the superficial dorsal horn of the spinal cord.
    Biotechnology and Biological Sciences Research Council
    2012 - 2014
     
  • Confocal microscopic studies in Neuroscience
    Wellcome Trust
    2010 - 2013
     
  • A quantitative description of glycinergic circuits in the ventral horn of the lumbar spinal cord.
    Wellcome Trust
    2009 - 2012
     
  • The role of spinal GluR4 containing AMPA receptors in pain mechanisms
    Tenovus-Scotland
    2007 - 2007
     
  • AMPA receptors in the spinal dorsal horn - synaptic distribution and plasticity in pain states
    Wellcome Trust
    2006 - 2010
     
  • P boutons in the rodent spinal cord
    Carnegie Trust for the Universities of Scotland
    2005 - 2006
     
  • Confocal microscopic studies in the spinal cord
    Wellcome Trust
    2002 - 2005
     
  • An Investigation of Spinal Cord Mechanisms Underlying Neuropathic Pain in Nerve Injury Models in the Rat
    Wellcome Trust
    2000 - 2006
     

Editorial Board

  • 2009 - ongoing: Frontiers in Neuroanatomy
  • 2004 - ongoing: Molecular Pain
  • 2000 - ongoing: Neuroscience

Grant Advisory Board

  • 2009 - ongoing: Wellcome Trust - Chairman of the External Advisory Board of the London Pain Consortium

Invited International Presentations

  • 2011: Hamburg, Germany - Refresher course, EFIC meeting, invited speaker
  • 2011: Heidleberg, Germany - EFIC Satellite symposium, speaker
  • 2011: Porto, Portugal - Departmental Seminar
  • 2009: Zurich, Switzerland - Invited seminar
  • 2009: Nagoya, Japan - Plenary lecturer, "Nagoya Pain 2009", (joint meeting of Japanese Association for the Study of Pain and Japanese Society of Pain Clinicians)
  • 2009: New York, USA - Plenary Lecture - World Institute of Pain (WIP) 5th World Congress
  • 2009: El Escorial, Spain - Invited speaker, London Pain Consortium summer school
  • 2009: Bordeaux, France - Symposium speaker, 9th Congress French Neuroscience Society
  • 2008: Glasgow, UK - Plenary Lecture - International Association for the Study of Pain (IASP) 12th World Congress
  • 2007: Basel, Switzerland - University of Basel
  • 2006: Lund, Sweden - Department of Experimental Medicine, Lund University
  • 2005: Xi'an, China - Invited talk: Fourth Military Medical University
  • 2005: La Jolla, CA, USA - Invited Seminar at Johnson and Johnson
  • 2005: Pecs, Hungary - 11th Annual Congress of the Hungarian Neuroscience Society (symposium speaker)
  • 2005: San Diego, USA - Mini Symposium, Department of Anesthesiology, University of California
  • 2004: Kyoto, Japan - International Federation of Associations of Anatomists (IFAA)
  • 2003: Torino, Italy - European Colloquium on Pain
  • 2003: Prague, Czech Republic - International Brain Research Organization (IBRO) Satellite symposium
  • 2002: Tubingen, Germany - German Physiological Society, Symposium speaker
  • 2002: San Diego, CA, USA - IASP 10th World Congress, Symposium speaker
  • 2001: Montreal, Canada - University of Montreal, Peptide Receptors.
  • 2001: Bristol, UK - Physiological Society, Symposium speaker

Professional Learned Society

  • 2011 - ongoing: Society of Biology - Fellow
  • 2006 - ongoing: Physiological Society - Member
  • 2001 - ongoing: International Association for the Study of Pain - Member
  • 1997 - ongoing: British Neuroscience Association - Member
  • 1997 - ongoing: Society for Neuroscience - Member
  • 1996 - ongoing: European Neuroscience Association - Member