Alison Lloyd Research Group

1985 - BSc Biochemistry - UCL
1989 - PhD - University of London (Chester Beatty Labs, Institute of Cancer Research)
Alison Lloyd
Tel: 020 7679 2240
Fax: 020 7679 7805
1984 Biochemistry Prize (University College London)
1989 EMBO Long Term Fellowship
1990 EEC Cancer Training Fellowship
1993 National Neurofibromatosis Foundation Young Investigator Award; Jaffa Fellowship
1998 Cancer Research UK Senior Cancer Research Fellowship
2004 Renewal of Cancer Research UK Senior Cancer Research Fellowship
2006 Readership in the Dept. of Biochemistry, UCL
2009 Professor of Cell Biology, FLS, UCL
Previous Posts: 
1989 - Postdoctoral Fellow - Institut de Chimie Biologique, Strasbourg, France
1993 - Postdoctoral Fellow - ICRF, London
1998 - CRUK Senior Cancer Research Fellow/Group Leader – LMCB, UCL
2009 - Professor of Cell Biology - UCL

Cell growth and tissue regeneration

Schwann cells are the main glial cells of the peripheral nervous system. Properties of these cells make them a powerful model system for studying two fundamental biological processes: (1) Cell growth (2) Tissue regeneration. Using a combination of primary in vitro culture systems and in vivo mouse models we aim to study the molecular and cellular mechanisms underlying the homeostatic regulation of these processes and how they become deregulated in diseases such as cancer. These studies should identify novel pathways important in these processes and new targets for regenerative medicine and cancer therapeutics.


Cell growth

Homeostasis in the adult requires a balance of positive and negative regulatory stimuli to control cell size and cell number. Whether a cell divides, grows or dies depends both on the levels of extracellular signals and the intrinsic ability of a cell to respond to these signals. Cell growth (addition of volume/mass), as cell proliferation and survival, requires an instructive signal but the signalling pathways and cell biology underlying cell growth are relatively understudied. Moreover, the additional mass required for tumour formation, requires deregulated growth pathways as well as a loss of proliferative controls but how oncogenic changes effect cell growth control is poorly understood.
Growth is often thought of as a uniform process however, specific cells contain different amounts of protein, membranes and organelles and these can change in response to specific stimuli. How cell set their size, count their organelles and adapt these to changing environments remain key unanswered questions in cell biology.


Enormous variation in cell size and organelle proportionality dependent on instructive extracellular signals.

Projects in the lab include the following

  • Identification of novel cell growth pathways
  • Regulation of organelle biogenesis
  • Cellular mechanisms underlying cell growth
  • Oncogenic regulation of cell growth

Tissue regeneration

The peripheral nervous system is one of the few tissues in the mammalian adult, which is capable of extensive regeneration. This process is all the more remarkable, in that repair can reconnect andre-establish fully transected nerves – requiring both the production of new tissue to bridge the gap between the nerve stumps and the accurate direction of regrowing axons back to their targets. Schwann cells are known to play a pivotal role in this process. In the adult, these highly specialised cells are normally in a quiescent state, myelinating larger axons or bundling together groups of smaller axons. Upon injury however, they dedifferentiate en masse to a progenitor/stem-like state and the proliferation and organisation of these cells is known to be critical for the repair process. Schwann cell number and state is strictly controlled by the axon both during development and following repair. Imbalances in this tightly regulated system would be predicted to result in either degenerative disorders or hyperproliferative disorders such as cancer. Consistent with this view, Schwann cell tumours, especially neurofibromas, resemble an unrepaired wounded nerve, in that Schwann cells within the tumours are dedifferentiated and proliferate in the absence of axonal contact in a mixture of fibroblasts and inflammatory cells.



Projects in the lab include the following

  • Understanding the plasticity of the Schwann cell differentiation state
  • Novel mouse models for studying tumour development in NF1
  • Role of the microenvironment in the repair process and cancer


Lab Members: 
Noelia Garcia Calavia
Anne-Laure Cattin
Melissa Collins
Xavier Garcia Fontana
Ilaria Napoli
Lucie Van Emmenis
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Number of items: 46.


Plotkin, SR; Albers, AC; Babovic-Vuksanovic, D; Blakeley, JO; Breakefield, XO; Dunn, CM; Evans, DG; (2014) Update from the 2013 International Neurofibromatosis Conference. Am J Med Genet A , 164A (12) 2969 - 2978. 10.1002/ajmg.a.36754.


Lloyd, AC; (2013) The regulation of cell size. Cell , 154 (6) 1194 - 1205. 10.1016/j.cell.2013.08.053.

Ribeiro, S; Napoli, I; White, IJ; Parrinello, S; Flanagan, AM; Suter, U; Parada, LF; (2013) Injury Signals Cooperate with Nf1 Loss to Relieve the Tumor-Suppressive Environment of Adult Peripheral Nerve. Cell Rep , 5 (1) pp. 126-136. 10.1016/j.celrep.2013.08.033. Green open access


Collins, MJ; Napoli, I; Ribeiro, S; Roberts, S; Lloyd, AC; (2012) Loss of Rb cooperates with Ras to drive oncogenic growth in mammalian cells. Curr Biol , 22 (19) 1765 - 1773. 10.1016/j.cub.2012.07.040.

Kalamarides, M; Acosta, MT; Babovic-Vuksanovic, D; Carpen, O; Cichowski, K; Evans, DG; Giancotti, F; (2012) Neurofibromatosis 2011: a report of the Children's Tumor Foundation annual meeting. Acta Neuropathol , 123 (3) 369 - 380. 10.1007/s00401-011-0905-0.

Napoli, I; Noon, LA; Ribeiro, S; Kerai, AP; Parrinello, S; Rosenberg, LH; Collins, MJ; (2012) A central role for the ERK-signaling pathway in controlling Schwann cell plasticity and peripheral nerve regeneration in vivo. Neuron , 73 (4) 729 - 742. 10.1016/j.neuron.2011.11.031.

Napoli, I; Noon, LA; Ribeiro, S; Kerai, AP; Parrinello, S; Rosenberg, LH; Collins, MJ; (2012) A central role for the ERK-signaling pathway in controlling Schwann cell plasticity and peripheral nerve regeneration in vivo. Neuron , 73 (4) 729 - 742. 10.1016/j.neuron.2011.11.031.

Roberts, SA; Lloyd, AC; (2012) Aspects of cell growth control illustrated by the Schwann cell. Curr Opin Cell Biol , 24 (6) 852 - 857. 10.1016/


Feber, A; Wilson, GA; Zhang, L; Presneau, N; Idowu, B; Down, TA; Rakyan, VK; (2011) Comparative methylome analysis of benign and malignant peripheral nerve sheath tumors. Genome Research , 21 (4) pp. 515-524. 10.1101/gr.109678.110. Gold open access


Danovi, D; Cremona, CA; Machado-da-Silva, G; Basu, S; Noon, LA; Parrinello, S; Lloyd, AC; (2010) A Genetic Screen for Anchorage-Independent Proliferation in Mammalian Cells Identifies a Membrane-Bound Neuregulin. PLOS ONE , 5 (7) , Article e11774. 10.1371/journal.pone.0011774. Green open access

Parrinello, S; Napoli, I; Ribeiro, S; Digby, PW; Fedorova, M; Parkinson, DB; Doddrell, RDS; (2010) EphB Signaling Directs Peripheral Nerve Regeneration through Sox2-Dependent Schwann Cell Sorting. CELL , 143 (1) 145 - 155. 10.1016/j.cell.2010.08.039.


Cremona, CA; Lloyd, AC; (2009) Loss of anchorage in checkpoint-deficient cells increases genomic instability and promotes oncogenic transformation. J CELL SCI , 122 (18) 3272 - 3281. 10.1242/jcs.047126.

Echave, P; Machado-da-Silva, G; Arkell, RS; Duchen, MR; Jacobson, J; Mitter, R; Lloyd, AC; (2009) Extracellular growth factors and mitogens cooperate to drive mitochondrial biogenesis. J CELL SCI , 122 (24) 4516 - 4525. 10.1242/jcs.049734. Gold open access

Parrinello, S; Lloyd, AC; (2009) Neurofibroma development in NF1-insights into tumour initiation. TRENDS CELL BIOL , 19 (8) 395 - 403. 10.1016/j.tcb.2009.05.003.


Parkinson, DB; Bhaskaran, A; Arthur-Farraj, P; Noon, LA; Woodhoo, A; Lloyd, AC; Feltri, ML; (2008) c-Jun is a negative regulator of myelination. J CELL BIOL , 181 (4) 625 - 637. 10.1083/jcb.200803013. Gold open access

Parrinello, S; Noon, LA; Harrisingh, MC; Digby, PW; Rosenberg, LH; Cremona, CA; Echave, P; (2008) NF1 loss disrupts Schwann cell-axonal interactions: a novel role for semaphorin 4F. GENE DEV , 22 (23) 3335 - 3348. 10.1101/gad.490608. Gold open access

Rubio, D; Garcia, S; De la Cueva, T; Paz, MF; Lloyd, AC; Bernad, A; Garcia-Castro, J; (2008) Human mesenchymal stem cell transformation is associated with a mesenchymal-epithelial transition. EXP CELL RES , 314 (4) 691 - 698. 10.1016/j.yexcr.2007.11.017.

Rubio, D; Garcia, S; Paz, MF; De la Cueva, T; Lopez-Fernandez, LA; Lloyd, AC; Garcia-Castro, J; (2008) Molecular Characterization of Spontaneous Mesenchymal Stem Cell Transformation. PLOS ONE , 3 (1) , Article e1398. 10.1371/journal.pone.0001398. Green open access


Echave, P; Conlon, IJ; Lloyd, AC; (2007) Cell size regulation in mammalian cells. CELL CYCLE , 6 (2) 218 - 224.

Noon, LA; Lloyd, AC; (2007) Treating leprosy: an Erb-al remedy? TRENDS PHARMACOL SCI , 28 (3) 103 - 105. 10.1016/


Lloyd, AC; (2006) Distinct functions for ERKs? J Biol , 5 (5) 13 - ?. 10.1186/jbiol46. Gold open access


Noon, LA; Lloyd, AC; (2005) Hijacking the ERK signaling pathway: Mycobacterium leprae shuns MEK to drive the proliferation of infected Schwann cells. Sci STKE , 2005 (309) pe52 - ?. 10.1126/stke.3092005pe52.

Rubio, D; Garcia-Castro, J; Martin, MC; de la Fuente, R; Cigudosa, JC; Lloyd, AC; Bernad, A; (2005) Spontaneous human adult stem cell transformation (Retracted article. See vol. 70, pg. 6682, 2010). CANCER RES , 65 (8) 3035 - 3039.


Conlon, I; Lloyd, AC; Raff, MC; (2004) Co-ordination of cell growth and cell-cycle progression in proliferating mammalian cells. In: Hall, MN and Raff, M and Thomas, G, (eds.) Cell Growth: Control of Cell Size. (85 - 101). Cold Spring Harbour Laboratory Press

Harrisingh, MC; Lloyd, AC; (2004) Ras/Raf/ERK signalling and NF1. CELL CYCLE , 3 (10) 1255 - 1258.

Harrisingh, MC; Lloyd, AC; (2004) Ras/Raf/ERK signalling and NF1: Implications for Neurofibroma Formation. Cell Cycle , 3 (10) 1255 - 1258.

Harrisingh, MC; Perez-Nadales, E; Parkinson, DB; Malcolm, DS; Mudge, AW; Lloyd, AC; (2004) The Ras/Raf/ERK signalling pathway drives Schwann cell dedifferentiation. EMBO J , 23 (15) 3061 - 3071. 10.1038/sj.emboj.7600309.

Perez-Nadales, E; Lloyd, AC; (2004) Essential function for ErbB3 in breast cancer proliferation. BREAST CANCER RES , 6 (3) 137 - 139. 10.1186/bcr792.


Mitchell, PJ; Perez-Nadales, E; Malcolm, DS; Lloyd, AC; (2003) Dissecting the contribution of p16(INK4A) and the Rb family to the Ras transformed phenotype. MOL CELL BIOL , 23 (7) 2530 - 2542. 10.1128/MCB.23.7.2530-2542.2003.


Lloyd, AC; (2002) Limits to lifespan. Nat Cell Biol , 4 (2) E25 - E27. 10.1038/ncb0202-e25.

Lloyd, AC; (2002) Limits to Lifespan. Nature Cell Biology , 4 0 - 0.


Mathon, NE; Lloyd, AC; (2001) Cell senescence and cancer. NAT REV CANCER , 1 (3) 203 - 213.

Mathon, NF; Malcolm, DS; Harrisingh, MC; Cheng, LL; Lloyd, AC; (2001) Lack of replicative senescence in normal rodent glia. SCIENCE , 291 (5505) 872 - 875.

Tang, DG; Tokumoto, YM; Apperly, JA; Lloyd, AC; Raff, MC; (2001) Lack of replicative senescence in cultured rat - Oligodendrocyte precursor cells. SCIENCE , 291 (5505) 868 - 871.


(2000) p53: Only ARF the story. Nature Cell Biology , 2 48 - 50.


Lloyd, AC; (1998) Ras versus cyclin-dependent kinase inhibitors. CURR OPIN GENET DEV , 8 (1) 43 - 48.


Lloyd, AC; Obermuller, F; Staddon, S; Barth, C; McMahon, M; Land, H; (1997) Co-operating Oncogenes Target Cyclin/Cdk Activity. Genes and Development , 11 663 - 677.

Sewing, A; Wiseman, B; Lloyd, AC; Land, H; (1997) high Intensity Raf Signal Causes Cell-Cycle Arrest Mediated by p21Cip1. Molecular and Cellular Biology , 17 5588 - 5597.


Lloyd, AC; Yancheva, N; Wasylyk, B; (1991) Transformation suppressor Activity of a Jun transcription Factor Lacking its Activation Domain. Nature , 352 635 - 638.


Flores, P; Gutman, A; Imler, JL; Lloyd, AC; Schneikert, J; Wasylyk, C; Wasylyk, B; (1989) AP1 and PEA3 are Nuclear targets for Transcriptional Activation by Non-nuclear oncogenes. Hormones and Cell Regulation , 198 57 - 62.

Lloyd, AC; Davies, SA; Crossley, I; Whitaker, M; Houslay, MD; Hall, A; Marshall, CJ; (1989) Bombesin stimulation of inositol 1,4,5-trisphosphate generation and intracellular calcium release is amplified in a cell line overexpressing the N-ras proto-oncogene. Biochemical Journal , 260 813 - 819.

Lloyd, AC; Paterson, HF; Morris, JD; Hall, A; Marshall, CJ; (1989) p21H-Ras Induced Morphological Transformation and Increases in c-myc Expression are Independent of functional Protein kinase C. The EMBO Journal , 8 1099 - 1104.

Marshall, CJ; Lloyd, AC; Morris, JD; Paterson, H; Price, B; Hall, A; (1989) Signal transduction by p21ras. International Journal of Cancer - Supplement , 4 29 - 31.

Morris, JDH; Price, B; Lloyd, AC; Self, AJ; Marshall, CJ; Hall, A; (1989) Scrape-loading of Swiss 3T3 cells with ras protein rapidly activates protein kinase C in the absence of phosphoinositide hydrolysis. Oncogene , 4 27 - 31.


Hall, A; Cales, C; Hancock, JF; Lloyd, AC; Self, A; Gardener, S; Houslay, MD; (1988) Analysis of mammalian Ras effector function. Cold Spring Harbor Symposium on Quantitative Biology , 53 855 - 862.


Lloyd, AC; Carpenter, CA; Saggerson, ED; (1986) Intertissue differences in the hysteretic behaviour of carnitine palmitoyltransferase in the presence of malonyl-CoA. Biochemical Journal , 237 289 - 291.

This list was generated on Thu Jul 23 11:13:00 2015 BST.