Xiu-Tang Cheng, Ning Huang, and Zu-Hang Sheng. (2022). Programming axonal mitochondrial maintenance and bioenergetics in neurodegeneration and regeneration.  Neuron  110, 1899-1923. PDF 

Joseph C. Roney, Xiu-Tang Cheng, and Zu-Hang Sheng. (2022). Neuronal endolysosomal transport and lysosomal functionality in maintaining axonostasis (Review).  Journal of Cell Biology   221, 3, 2022. PDF

Sunan Li and Zu-Hang Sheng. (2022). Energy matters: presynaptic metabolism and the maintenance of synaptic transmission.  Nature Reviews Neuroscience  23, 4-22. PDF


Kelly A. Chamberlain, Ning Huang (co-first author), Yuxiang Xie, Francesca LiCausi, Sunan Li, Yan Li, and Zu-Hang Sheng. (2021). Oligodendrocytes enhance axonal energy metabolism by deacetylation of mitochondrial proteins through transcellular delivery of SIRT2.  Neuron 109, 3456-3472. PDF

  • NINDS/NIH Press: Signaling from neighboring cells provides power boost within axons. PDF
  • Neuron Preview by Eva-Maria Albers: Superfood for axons: Glial exosomes boost axonal energetics by delivery of SIRT2.  Neuron  109, 3397-3400, 2021. PDF

Joseph C Roney, Sunan Li, Tamar Farfel-Becker, Ning Huang, Tao Sun, Yuxiang Xie, Xiu-Tang Cheng, Mei-Yao Lin, Frances M Platt, and Zu-Hang Sheng. (2021). Lipid-mediated impairment of axonal lysosome transport contributing to autophagic stress.  Autophagy  17, 1796-1798. PDF

Ning Huang, Sunan Li, Yuxiang Xie, Qi Han, Xiao-Ming Xu, and Zu-Hang Sheng. (2021). Reprogramming an energetic AKT-PAK5 axis boosts axon energy supply and facilitates neuron survival and regeneration after injury and ischemia.  Current Biology   31, 3098-3114. PDF

  • Current Biology | Dispatch by Twiss et al.,Neurobiology: Resetting the axon’s batteries”. PDF

Xiu-Tang Cheng and Zu-Hang Sheng. (2021). Neurobiology: A Pathogenic Tug of War.  Current Biology  31, 491–493. PDF

Joseph C Roney, Sunan Li*, Tamar Farfel-Becker*, Ning Huang, Tao Sun, Yuxiang Xie, Xiu-Tang Cheng, Mei-Yao Lin, Frances M Platt, and Zu-Hang Sheng. (2021)Lipid-mediated motor-adaptor sequestration impairs axonal lysosome delivery leading to autophagic stress and dystrophy in Niemann-Pick type C.  Developmental Cell  56, 1452-1468. PDF

  • Developmental Cell Preview by Yap and Winckler "Lysosomes to the rescue: Anterograde axonal lysosome transport and neuronal proteostasisPDF
  • Nature Reviews Molecular Cell Biology RESEARCH HIGHLIGHT "Lysosome transport interrupted".  PDF

Gui-Jing Xiong, Xiu-Tang Cheng, Tao Sun, Yuxiang Xie, Ning Huang, Sunan Li, Meo-Yao Lin, and Zu-Hang Sheng. (2021). Defective axonal transport associates with autism-like synaptic dysfunction and social behavioral traits.  Molecular Psychiatry  26, 1472-1490.  PDF


Sunan Li, Gui-Jing Xiong, Ning Huang, and Zu-Hang Sheng. (2020). The crosstalk of energy sensing and mitochondrial anchoring sustains synaptic efficacy by maintaining presynaptic metabolism.  Nature Metabolism  2, 1077-1095.  PDF

  • NIH Press: "NIH scientists reveal how the brain may fuel intense neural communication"PDF
  • Science Research Highlight "Mitochondrial anchoring in synapses". PDF

Xiu-Tang Cheng and Zu-Hang Sheng. (2020). Developmental regulation of microtubule-based axonal mitochondria trafficking and anchoring in health and diseases. Developmental Neurobiology  81, 284-299  (Review). PDF

Qi Han, Yuxiang Xie, Josue D Ordaz, Andrew J Huh, Ning Huang, Wei Wu, Naikui Liu, Kelly A Chamberlain, Zu-Hang Sheng (lead corresponding), Xiao-Ming Xu (2020). Recovering energy deficits promotes CNS axonal regeneration and functional restoration after spinal cord injuryCell Metabolism  31, 623-641.  PDF

  • NIH Press: "Boosting energy levels within damaged nerves may help them heal".  PDF

Rajat Puri, Xiu-Tang Cheng, Mei-Yao Lin, Ning Huang, and Zu-Hang Sheng (2019). Defending stressed mitochondria: uncovering the role of MUL1 in suppressing neuronal mitophagy.  Autophagy  16, 176-178. PDF

Tamar Farfel-Becker, Joseph V. Roney, Xiu-Tang Cheng, Sunan Li, Sean R. Cuddy, and Zu-Hang Sheng. (2020). The secret life of degradative lysosomes in axons: delivery from the soma, enzymatic activity, and local autophagic clearance.  Autophagy  16, 167-168. PDF

Dinesh C Joshi, Chuan-Li Zhang, Lavanya Babujee, Jason D Vevea, Benjamin K August, Zu-Hang Sheng, Edwin R Chapman, Timothy M Gomez, Shing Yan Chiu (2019). Inappropriate Intrusion of an Axonal Mitochondrial Anchor into Dendrites Causes Neurodegeneration. Cell Reports  29, 685-696. PDF

Rajat Puri, Xiu-Tang Cheng, Mei-Yao Lin, Ning Huang, and Zu-Hang Sheng (2019). Mul1 restrains Parkin-mediated mitophagy in mature neurons by maintaining ER-mitochondrial contacts.  Nature Communications  10, 3645. PDF

Tamar Farfel-Becker, Joseph V. Roney, Xiu-Tang Cheng, Sunan Li, Sean R. Cuddy, and Zu-Hang Sheng (2019). Neuronal soma-derived degradative lysosomes are continuously delivered to distal axons to maintain local degradation capacity.  Cell Reports  28, 51-64. PDF

Kelly A. Chamberlain and Zu-Hang Sheng (2019). Mechanisms for the maintenance and regulation of axonal energy supply. Journal of Neuroscience Research 97, 897-913 (Review). PDF


Xiu-Tang Cheng, Yuxiang Xie, Bing Zhou, Ning Huang, Tamar Farfel-Becker, Zu-Hang Sheng. (2018). Revisiting LAMP1 as a marker for degradative autophagy-lysosomal organelles in the nervous system.  Autophagy. 14, 1472-1474. PDF

Xiu-Tang Cheng,  Yuxiang Xie,  Bing Zhou, Ning Huang, Tamar Farfel-Becker, and Zu-Hang Sheng (2018). Characterization of LAMP1-labeled non-degradative lysosomal and endocytic compartments in nervous systems.  Journal of Cell Biology  217, 3127-3139.  PDF

  • Featured in the JCB Spotlight Article by Kulkarni and Maday “Neuronal endosomes to lysosomes: A journey to the soma”  PDF

Mei-Yao Lin, Xiu-Tang Cheng (co-first author), Prasad Tammineni, Yuxiang Xie, Bing Zhou, Qian Cai, Zu-Hang Sheng (2017)Releasing Syntaphilin Removes Stressed Mitochondria from Axons Independent of Mitophagy under pathophysiological Conditions.  Neuron  94, 595-610. PDF

Zu-Hang Sheng (2017), The Interplay of Axonal Energy Homeostasis and Mitochondrial Trafficking and Anchoring. (Invited Review), Trends in Cell Biology. 27, 403-416.  PDF


Bing Zhou, Panpan Yu, Mei-Yao Lin, Tao Sun, Yanmin Cheng and Zu-Hang Sheng (2016). Facilitation of axon regeneration by enhancing mitochondrial transport and rescuing energy deficits.  Journal of Cell Biology. 204, 103-119.  PDF

  • Rockefeller Press: "Mobilizing mitochondria may be key to regenerating damaged neuronsPDF
  • NATURE | RESEARCH HIGHLIGHT "Mitochondria make nerves grow".  PDF
  • The New England Journal of Medicine RESEARCH HIGHLIGHT "Mitochondrial mobility and neuronal recovery".  PDF

Natalia S. Morsci, David H. Hall, Monica Driscoll, and Zu-Hang Sheng (2016). Age-related phasic patterns of mitochondrial maintenance in adult C. elegans neuronsJournal of Neuroscience, 36, 1373-1385.  PDF


Yuxiang Xie, Bing Zhou (co-first author), Mei-Yao Lin, Shiwei Wang, Kevin D. Foust, and Zu-Hang Sheng (2015). Endolysosomal Deficits Augment Mitochondria Pathology in Spinal Motor Neurons of Asymptomatic fALS Mice.   Neuron  87, 355-370.  PDF

  • NIH Press: “Neurons’ broken machinery piles up in ALS: NIH scientists identify a transport defect in a model of familial ALS”.  PDF

Jerome Di Giovanni & Zu-Hang Sheng (2015)Regulation of synaptic activity by snapin-mediated endolysosomal transport and sorting.  The EMBO Journal, 34, 2059-2077.  PDF

Xiu-Tang Cheng, Bing Zhou, Mei-Yao Lin, Qian Cai, and Zu-Hang Sheng (2015). Axonal autophagosomes recruit dynein for retrograde transport. through fusion with late endosomes. Journal of Cell Biology.  209, 377-386.  PDF

Mei-Yao Lin and Zu-Hang Sheng (2015). Regulation of mitochondrial transport in neurons (Review). Exp Cell Res, 334, 35-44.  PDF

Dinesh C. Joshi, Chuan-Li Zhang, Tien-Min Lin, Anchal Gusain, Melissa G. Harris, Esther Tree, Yewin Yin, Connie Wu, Zu-Hang Sheng, Robert J Dempsey, Zsuzsanna Fabry, and Shing Yan Chiu (2015). Deletion of Mitochondrial Anchoring Protects Dysmyelinating Shiverer: Implications for Progressive MS. Journal of Neuroscience, 35, 5293-5306. 


Zu-Hang Sheng (2014). Mitochondrial trafficking and anchoring in neurons: New insight and implications. (Invited Review), Journal of Cell Biology, 204, 1087-98.  PDF

Yun J, Puri R, Yang H, Lizzio MA, Wu C, Sheng ZH, Guo M (2014). MUL1 acts in parallel to the PINK1/parkin pathway in regulating mitofusin and compensates for loss of PINK1/parkin.  eLife. 3, e01958. PDF

Nobuhiko Ohno, Hao Chiang, Don J Mahad, Grahame Kidd, Liping Liu, Richard M. Ransohoff, Zu-Hang Sheng, Hitoshi Komuro and Bruce D. Trapp (2014). Mitochondrial immobilization mediated by syntaphilin facilitates survival of demyelinated axonsPNAS, 111, 9953-9958.  PDF


Yanmin Chen and Zu-Hang Sheng (2013). Kinesin-1-syntaphilin coupling mediates activity-dependent regulation of axonal mitochondrial transport.  Journal of Cell Biology  202, 351-364.  PDF

  • Featured in JCB by an In Focus Article: “Syntaphilin puts the brakes on axonal mitochondriaPDF

Tao Sun, Haifa Qiao (co-first author), Ping-Yue Pan, Yanming Chen, and Zu-Hang Sheng (2013)Mobile axonal mitochondria contribute to the variability of presynaptic strength.  Cell Reports.  4, 413-419. PDF

  • NIH Press "NIH researchers discover how brain cells change their tune"PDF 

Qian Cai, Hesham M Zakaria, and Zu-Hang Sheng (2012). Long time-lapse imaging reveals unique features of PARK2/Parkin-mediated mitophagy in mature cortical neurons.  Autophagy  8, 976-978.  PDF

Bing Zhou, Qian Cai, Yuxiang Xie, and Zu-Hang Sheng (2012). Snapin recruits dynein to BDNF-TrkB signaling endosomes for retrograde axonal transport and is essential for dendrite growth of cortical neuronsCell Reports  2, 42-51.  PDF

Kim HJ, Zhong Q, Zu-Hang Sheng, Yoshimori T, Liang C, Jung JU (2012). Beclin 1-interacting autophagy protein Atg14L targets SNARE-associated protein Snapin to coordinate endocytic traffickingJournal of Cell Science, 125, 4740-4750.  

Qian Cai, Hesham Mostafa Zakaria, Anthony Simone, and Zu-Hang Sheng (2012). Spatial Parkin Translocation and Degradation of Depolarized Mitochondria via Mitophagy in Live Cortical Neurons.  Current Biology  22, 545-552.  PDF

Zu-Hang Sheng & Qian Cai (2012)Mitochondrial Transport in Neurons: Impact on Synaptic Homeostasis and Neurodegeneration (Invited Review). Nature Reviews Neuroscience, 13, 77-93.  PDF


Qian Cai and Zu-Hang Sheng (2011). Uncovering the role of Snapin in regulating autophagy-lysosomal function. Autophagy  7, 445-447. PDF

Bing Zhou, Yi-Bing Zhu, Lin Lin, Qian Cai, and Zu-Hang Sheng (2011). Snapin deficiency is associated with developmental defects of the central nervous system, Bioscience Reports, 31, 151-158.

Qian Cai, Matthew Davis, and Zu-Hang Sheng (2011). Regulation of Axonal Mitochondrial Transport and Its Impact on Synaptic Transmission. Neuroscience Research, 70, 9-15 (invited review)

Yi-Bing Zhu, Zu-Hang Sheng (2011). Increased Axonal Mitochondrial Mobility Does Not Slow ALS-like Disease in Mutant SOD1 Mice. The Journal of Biological Chemistry, 286, 23432-23440. PDF


Qian Cai, Li Lu, Jin-Hua Tian, Yi-Bing Zhu, Haifa Qiao, and Zu-Hang Sheng (2010)Snapin-regulated late endosomal transport is critical for efficient autophagy-lysosomal function in neurons.  Neuron  68, 73-86.  PDF

  • Neuron Preview by Yuzaki:  “Snapin snaps into the dynein complex for late endosome-lysosome trafficking and autophagy".  PDF

Qian Cai and Zu-Hang Sheng (2009)Moving or Stopping Mitochondria: Miro as a Traffic Cop by Sensing Calcium.  Neuron  61, 493-496.  PDF

Qian Cai and Zu-Hang Sheng (2009). Molecular Motors and Synaptic Assembly. The Neuroscientists, 15, 78-89.  PDF

Huan Ma, Qian Cai, Wenbo Lu, Zu-Hang Sheng (co-corresponding author), Sumiko Mochida (2009). KIF5B Motor Adaptor Syntabulin Maintains Synaptic Transmission in Sympathetic NeuronsJournal of Neuroscience, 29, 13019-13029

Qian Cai and Zu-Hang Sheng (2009). Mitochondrial Transport and Docking in AxonsExperimental Neurology, 218, 257-267

Ping-Yue Pan, Jin-Hua Tian, Zu-Hang Sheng (2009). Snapin facilitates the synchronization of synaptic vesicle fusion. Neuron, 61, 412-424.  PDF

Yan-Min Chen, Claudia Gerwin, and Zu-Hang Sheng (2009). Dynein Light Chain LC8 Regulates Syntaphilin-Mediated Mitochondrial Docking in AxonsJournal of Neuroscience, 29, 9429-9438


Jian-Sheng Kang, Jin-Hua Tian*, Ping-Yue Pan*, Philip Zald, Cuiling Li, Chuxia Deng, and Zu-Hang Sheng (2008). Docking of Axonal Mitochondria by Syntaphilin Controls their Mobility and Affects Short-term Facilitation (*equal contribution).  Cell  132, 137-248.  PDF

  • Nature Reviews Neuroscience | RESEARCH HIGHLIGHT  “Mitochondria in the dock".  PDF

AG Miriam Leenders, Lin Lin, Li-Dong Huang, Claudia Gerwin, Pei-Hua Lu, and Zu-Hang Sheng (2008). The Role of MAP1A Light Chain 2 in Synaptic Surface Retention of Cav2.2 Channels in Hippocampal NeuronsJournal of Neuroscience, 28, 11333-11346

Wenbo Lu, Huan Ma, Zu-Hang Sheng, and Sumiko Mochida. (2008). Dynamin and activity regulate synaptic vesicle recycling in sympathetic neurons. The Journal of Biological Chemistry, 284, 1930-1937


Qain Cai, Ping-Yue Pan, and Zu-Hang Sheng (2007). Syntabulin-kinesin-1 family 5B-mediated axonal transport contributes to activity-dependent presynaptic assembly. Journal of Neuroscience, 27, 7284-7296.  PDF


Ping-Yue Pan, Qian Cai, Lin Lin, Pei-Hua Lu, Shu-Min Duan, and Zu-Hang Sheng. (2005). SNAP29-mediated modulation of synaptic transmission in cultured hippocampal neuronsThe Journal of Biological Chemistry, 280, 25769-25779

Qian Cai, Claudia Gerwin, and Zu-Hang Sheng (2005). Syntabulin-mediated anterograde transport of mitochondria along the neuronal processesJournal of Cell Biology, 170, 959-969

Jin-Hua Tian, Zheng-Xing Wu, Michael Unzicker, Li Lu, Qian Cai, Cuiling Li, Claudia Schirra, Ulf Matti, David Stevens, Chuxia Deng, Jens Rettig, and Zu-Hang Sheng (2005). The Role of Snapin in Neurosecretion: Snapin Knockout Mice Exhibit Impaired Calcium-dependent Exocytosis of Large Dense-core Vesicles in Chromaffin CellsJournal of Neuroscience, 25, 10546-10555 (With Weekly Editorial News)

Xiao-Ke Chen, Lie-Cheng Wang, Yang Zhou, Qian Cai, Murali Prakriya, Kai-Lai Duan, Zu-Hang Sheng, Christopher Lingle & Zhuan Zhou (2005). Activation of GPCRs modulates quantal size in chromaffin cells through Gbg and PKC. Nature Neuroscience, 8, 1160-1168

Wu LJ, Leenders AG, Cooperman S, Meyron-Holtz E, Smith S, Land W, Tsai RY, Berger UV, Sheng ZH, Rouault TA (2004). Expression of the iron transporter ferroportin in synaptic vesicles and the blood-brain barrierBrain Res. , 1001, 108-17

Pratima Thakur, David R. Stevens, Zu-Hang Sheng and Jens Rettig (2004). Effects of PKA-mediated phosphorylation of Snapin on synaptic transmission in cultured hippocampal neuronsJournal of Neuroscience, 24, 6476-6481

Qingning Su, Qian Cai (co-first author), Claudia Gerwin, Carolyn L. Smith, Zu-Hang Sheng (2004). Syntabulin: a microtubule-associated protein implicated in syntaxin trafficking in neurons. Nature Cell Biology, 6, 941-953. 

Judit Boczan, A. G. Miriam Leenders, and Zu-Hang Sheng (2004). Phosphorylation of syntaphilin by PKA modulates its interaction with syntaxin-1 and annuls its inhibitory effect on vesicle exocytosisThe Journal of Biological Chemistry, 279, 18911-18919.

Brij B. Singh, Timothy P. Lockwich, Bidhan C. Bandyopadhyay, Xibao Liu, Sunitha Bollimuntha, So-ching Brazer, Christian Combs, Sunit Das, A.G. Miriam Leenders, Zu-Hang Sheng, Mark A. Knepper, Suresh V. Ambudkar, and Indu S. Ambudkar (2004). VAMP2-Dependent Exocytosis Regulates Plasma Membrane Insertion of TRPC3 Channels and Contributes to Agonist-Stimulated Ca2+ Influx. Molecular Cell, 15, 635-646.

Haitao Song, Liping Nie, Adrian Rodriguez-Contreras, Zu-Hang Sheng, and Ebenexer Yamoah (2003). Functional interaction of auxiliary subunits and synaptic proteins with Ca1.3 may impart hair cell Ca current properties. J Neurophysiol., 89, 1143.

Jin-Hua Tian, Sunit Das, and Zu-Hang Sheng (2003). Ca-dependent phosphorylation of syntaxin-1A by DAP-kinase regulates its interaction with Munc-18The Journal of Biological Chemistry, 278, 26265-26274.

Sunit Das, Judit Boczan, Claudia Gerwin, Philip B. Zald and Zu-Hang Sheng (2003). Regional and developmental regulation of syntaphilin expression in the brain: a candidate molecular element of synaptic functional differentiationMolecular Brain Research, 116, 38-49.

Sunit Das, Claudia Gerwin, and Zu-Hang Sheng (2003). Syntaphilin binds to dynamin-1 and inhibits dynamin-dependent endocytosis. The Journal of Biological Chemistry, 278, 41221-41226.

Milan G. Chheda, Uri Ashery, Pratima Thakur, Jens Rettig,and Zu-Hang Sheng (2001). PKA phosphorylation of Snapin: modulating its interaction with the SNARE complexNature Cell Biology, 3, 331-338.

Qingning Su, Sumiko Mochida, Jin-Hua Tian, Rashi Mehta,and Zu-Hang Sheng (2001). SNAP-29: A General SNARE Protein that Inhibits SNARE Disassembly and is Implicated in Synaptic TransmissionProc. Natl. Acad. Sci. USA, 98, 1438-1443.

Guifang Lao, Volker Scheuss, Claudia M. Gerwin, Qingning Su, Sumiko Mochida, Jens Rettig, and Zu-Hang Sheng. (2000). Syntaphilin: a syntaxin-1 clamp that controls SNARE assemblyNeuron, 25, 191-201.

Jeffrey M. Ilardi, Sumiko Mochida, and Zu-Hang Sheng (1999). Snapin: a SNARE-associated protein implicated in synaptic transmissionNature Neuroscience, 2, 119-124.

Lucas D. Pozzo-Miller, Wolfram Gottschalk, Li Zhang, Kathryn McDermott, Chikara Oho, Zu-Hang Sheng, and Bai Lu (1999). The role of BDNF in Hippocampal synaptic plasticity: impairment in synaptic vesicle docking and synaptic protein distribution in BDNF knockout mice. J. Neuroscience, 19, 4972-4983.

Zu-Hang Sheng, Ruth E. Westenbroek, and William A. Catterall (1998). Physical link and functional coupling of presynaptic calcium channels and synaptic vesicle docking/fusion machinery. J. Bioenergetics & Biomembranes, 30, 335-345.

Zu-Hang Sheng, Charles Yokoyoma, and William A. Catterall (1997). Interaction of the synprint site of N-type calcium channels with the C2B domain of synaptotagmin I. Proc. Natl. Acad. Sci. USA, 94, 5405-5410.

Jens Rettig, Christian Heinemann, Uri Ashery, Zu-Hang Sheng, Charles T. Yokoyama, William A. Catterall, and Erwin Neher (1997). Alteration of Ca2+ dependence of neurotransmitter release by disruption of Ca2+ channel/SNARE protein interactionJ Neuroscience, 17, 6647-6656.

Charles Yokoyama, Zu-Hang Sheng, and William A. Catterall (1997). Phosphorylation of the synprint site on N-type calcium channels inhibits interactions with SNARE proteinJ Neuroscience, 17, 6929-6938.

Sumiko Mochida, Zu-Hang Sheng, Carl Baker, Haruo Kobayashi, and William A. Catterall (1996). Inhibition of neurotransmission by peptides containing the synaptic protein interaction site of N-type calcium channelsNeuron, 17, 781-788.

Jens Rettig, Zu-Hang Sheng, D. Kyle Kim, Connie D. Hodson, Terry P. Snutch, and William A. Catterall (1996). Isoform-specific interaction of the a1A subunits of brain calcium channels with the presynaptic proteins syntaxin and SNAP-25. Proc. Natl. Acad. Sci. USA, 93, 7363-7368.

Zu-Hang Sheng, Jens Rettig, Terry Cook, and William A. Catterall (1996). Calcium dependent interaction of neuronal N-type calcium channels with presynaptic fusion proteins. Nature, 379, 451-454.

Zu-Hang Sheng, Jens Rettig, Masami Takahashi, and William A. Catterall (1994). Identification of a syntaxin-binding site on N-type calcium channels. Neuron, 13, 1303-1313.

Roland G. Kallen, Zu-Hang Sheng, Liquiong Chen, Jan Yang, Rogart, R.B., and Robert L. Barchi (1990). Primary structure and differential expression of a sodium channel characteristic of immature and denervated rat skeletal muscle. Neuron, 4, 233-242.

Trimmer, J.S., Cooperman, S.S., Tomiko, S.A., Zhou, J., Crean, S.M., Boyle, M.B., Kallen, R.G., Zu-Hang Sheng, Barchi, R.L., Sigworth, F.J., Goodman, R.G., Agnew, S.A., and Mandel, G (1989). Primary structure and functional expression of a mammalian skeletal muscle sodium channel. Neuron, 3, 33-49.