Abstract
This chapter explores the current understanding of the hormonal regulation of auditory function in songbirds by focusing on three themes. The first section is an overview of seasonal changes in the auditory pathway that are regulated by hormones. Next, the concept of the songbird brain as both a source and a target of neuromodulatory steroid hormones is discussed in the context of auditory function. Finally, the way that hormones interact with classical neurotransmitter systems (the biogenic amines dopamine, norepinephrine, and serotonin) to modulate auditory processing is presented. Reflecting on the sum total of these studies, understanding of the hormonal regulation of auditory function in songbirds has progressed considerably in the past few decades. More broadly, the field of songbird neuroethology has been continually propelled by an integrative perspective that examines the development, evolution, and hormonal modulation of neural circuits for song production, learning, and processing. This holistic approach to songbird neuroethology research, inspired by Niko Tinbergen and Peter Marler, will continue to be important as an increasing number of tools become available to explore the brain and behavior of songbirds.
This is a preview of subscription content, log in via an institution.
Buying options
Tax calculation will be finalised at checkout
Purchases are for personal use only
Learn about institutional subscriptionsReferences
Abizaid A, Mezei G, Horvath TL (2004) Estradiol enhances light-induced expression of transcription factors in the SCN. Brain Res 1010(1–2):35–44
Acharya KD, Veney SL (2012) Characterization of the G-protein-coupled membrane-bound estrogen receptor GPR30 in the zebra finch brain reveals a sex difference in gene and protein expression. Dev Neurobiol 72(11):1433–1446
Adkins-Regan E (2005) Hormones and animal social behavior. Princeton University Press, Princeton
Alward BA, Balthazart J, Ball GF (2013) Differential effects of global versus local testosterone on singing behavior and its underlying neural substrate. Proc Natl Acad Sci U S A 110(48):19573–19578
Arch VS, Narins PM (2009) Sexual hearing: the influence of sex hormones on acoustic communication in frogs. Hear Res 252(1–2):15–20
Arnold AP, Nottebohm F, Pfaff DW (1976) Hormone concentrating cells in vocal control and other areas of brain of zebra finch (Poephila-Guttata). J Comp Neurol 165(4):487–511
Ball GF, Riters LV, Balthazart J (2002) Neuroendocrinology of song behavior and avian brain plasticity: multiple sites of action of sex steroid hormones. Front Neuroendocrinol 23(2):137–178
Ball, G. F., Castelino, C. B., Maney, D. L., Appeltants, D., & Balthazart, J. (2003). The activation of birdsong by testosterone - Multiple sites of action and role of ascending catecholamine projections Steroids and the Nervous System, 1007, 211–231
Balthazart J, Ball GF (2006) Is brain estradiol a hormone or a neurotransmitter? Trends in Neurosci 29(5):241–249
Balthazart J, Ball GF (eds) (2012) Brain aromatase, estrogens, and behavior. Oxford University Press, Oxford
Balthazart J, Choleris E, Remage-Healey L (2018) Steroids and the brain: 50years of research, conceptual shifts and the ascent of non-classical and membrane-initiated actions. Horm Behav 99:1–8
Barclay SR, Harding CF (1988) Androstenedione modulation of monoamine levels and turnover in hypothalamic and vocal control nuclei in the male zebra finch: steroid effects on brain monoamines. Brain Res 459(2):333–343
Barclay SR, Harding CF (1990) Differential modulation of monoamine levels and turnover rates by estrogen and/or androgen in hypothalamic and vocal control nuclei of male zebra finches. Brain Res 523(2):251–262
Bentley G (2000) Stimulatory effects on the reproductive axis in female songbirds by conspecific and heterospecific male song. Horm Behav 37(3):179–189
Bernard DJ, Bentley GE, Balthazart J, Turek FW, Ball GF (1999) Androgen receptor, estrogen receptor alpha, and estrogen receptor beta show distinct patterns of expression in forebrain song control nuclei of European starlings. Endocrinol 140(10):4633–4643
Boyd SK, Moore FL (1990) Evidence for gaba involvement in stress-induced inhibition of male amphibian sexual-behavior. Horm Behav 24(1):128–138
Brenowitz EA (2004) Plasticity of the adult avian song control system. Behavioral Neurobiology of Birdsong 1016:560–585
Brenowitz EA, Lent K (2002) Act locally and think globally: intracerebral testosterone implants induce seasonal-like growth of adult avian song control circuits. Proc Natl Acad Sci 99(19):12421–12426
Brenowitz EA, Nalls B, Wingfield JC, Kroodsma DE (1991) Seasonal-changes in avian song nuclei without seasonal-changes in song repertoire. J Neurosci 11(5):1367–1374
Caras ML, Remage-Healey L (2016) Modulation of peripheral and central auditory processing by estrogens in birds. In: Bass AH (ed) Hearing and hormones, vol 57. Springer, Heidelberg, pp 77–100
Caras ML, Brenowitz E, Rubel EW (2010) Peripheral auditory processing changes seasonally in Gambel’s white-crowned sparrow. J Comp Physiol A Neuroethol Sens Neural Behav Physiol 196(8):581–599
Caras ML, O’Brien M, Brenowitz EA, Rubel EW (2012) Estradiol selectively enhances auditory function in avian forebrain neurons. J Neurosci 32(49):17597–17611
Caras ML, Sen K, Rubel EW, Brenowitz EA (2015) Seasonal plasticity of precise spike timing in the avian auditory system. J Neurosci 35(8):3431–3445
Cardin JA, Schmidt MF (2004) Noradrenergic inputs mediate state dependence of auditory responses in the avian song system. J Neurosci 24(35):7745–7753
Castelino CB, Schmidt MF (2010) What birdsong can teach us about the central noradrenergic system. J Chem Neuroanat 39(2):96–111
Chakraborty, M., & Burmeister, S. S. (2015). Effects of estradiol on neural responses to social signals in female tungara frogs. J Exp Biol, 218(Pt 22), 3671–3677
Chao A, Paon A, Remage-Healey L (2014) Dynamic variation in forebrain estradiol levels during song learning. Dev Neurobiol
Chew SJ, Mello C, Nottebohm F, Jarvis E, Vicario DS (1995) Decrements in auditory responses to a repeated conspecific song are long-lasting and require 2 periods of protein-synthesis in the songbird forebrain. Proc Natl Acad Sci 92(8):3406–3410
Crimins JL, Wang AC, Yuk F, Puri R, Janssen WGM, Hara Y, Rapp PR, Morrison JH (2017) Diverse synaptic distributions of G protein-coupled estrogen receptor 1 in monkey prefrontal cortex with aging and menopause. Cereb Cortex 27(3):2022–2033
De Groof G, Poirier C, George I, Hausberger M, Van der Linden A (2013) Functional changes between seasons in the male songbird auditory forebrain. Front Behav Neurosci 7:196
De Groof G, Balthazart J, Cornil CA, Van der Linden A (2017) Topography and lateralized effect of acute aromatase inhibition on auditory processing in a seasonal songbird. J Neurosci 37(16):4243–4254
Del Negro C, Edeline JM (2002) Sex and season influence the proportion of thin spike cells in the canary HVc. Neuroreport 13(16):2005–2009
Del Negro C, Lehongre K, Edeline JM (2005) Selectivity of canary HVC neurons for the bird’s own song: modulation by photoperiodic conditions. J Neurosci 25(20):4952–4963
Foradori C, Weiser M, Handa R (2007) Non-genomic actions of androgens. Front Neuroendocrinol
Forlano PM, Deitcher DL, Myers DA, Bass AH (2001) Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source. J Neurosci 21(22):8943–8955
Forlano PM, Schlinger BA, Bass AH (2006) Brain aromatase: new lessons from non-mammalian model systems. Front Neuroendocrinol 27(3):247–274
Fusani L, Van’t Hof T, Hutchison JB, Gahr M (2000) Seasonal expression of androgen receptors, estrogen receptors, and aromatase in the canary brain in relation to circulating androgens and estrogens. J Neurobiol 43(3):254–268
Gall MD, Salameh TS, Lucas JR (2013) Songbird frequency selectivity and temporal resolution vary with sex and season. Proc Biol Sci 280(1751):20122296
Gentner TQ, Hulse SH (2000) Female European starling preference and choice for variation in conspecific male song. Anim Behav 59(2):443–458
Gentner TQ, Hulse SH, Duffy D, Ball GF (2001) Response biases in auditory forebrain regions of female songbirds following exposure to sexually relevant variation in male song. J Neurobiol 46(1):48–58
Gobes SMH, Bolhuis JJ (2007) Birdsong memory: a neural dissociation between song recognition and production. Curr Biol 17(9):789–793
Heimovics SA, Ferris JK, Soma KK (2015) Non-invasive administration of 17beta-estradiol rapidly increases aggressive behavior in non-breeding, but not breeding, male song sparrows. Horm Behav 69:31–38
Henry KS, Lucas JR (2009) Vocally correlated seasonal auditory variation in the house sparrow (Passer domesticus). J Exp Biol 212(23):3817–3822
Hofmann HA, Renn SC, Rubenstein DR (2016) Introduction to symposium: new frontiers in the integrative study of animal behavior: nothing in neuroscience makes sense except in the light of behavior. Int Comp Biol 56(6):1192–1196
Holveck MJ, Riebel K (2007) Preferred songs predict preferred males: consistency and repeatability of zebra finch females across three test contexts. Anim Behav 74:297–309
Ikeda M, Rensel MA, Schlinger BA, Remage-Healey L (2014) In vivo detection of fluctuating brain steroid levels in zebra finches. Cold Spring Harb Protoc
Ikeda MZ, Jeon SD, Cowell RA, Remage-Healey L (2015) Norepinephrine modulates coding of complex vocalizations in the songbird auditory cortex independent of local neuroestrogen synthesis. J Neurosci 35(25):9356–9368
Kabelik D, Schrock SE, Ayres LC, Goodson JL (2011) Estrogenic regulation of dopaminergic neurons in the opportunistically breeding zebra finch. Gen Comp Endocrinol 173(1):96–104
Kelly MJ, Moss RL, Dudley CA (1976) Differential sensitivity of preoptic-septal neurons to microelectrophoresed estrogen during the estrous cycle. Brain Res 114(1):152–157
Krentzel AA, Macedo-Lima M, Ikeda MZ, Remage-Healey L (2018) A membrane G-protein coupled estrogen receptor is necessary but not sufficient for sex-differences in zebra finch auditory coding. Endocrinology 159(3):1360–1376
Lattin CR, Stabile FA, Carson RE (2017) Estradiol modulates neural response to conspecific and heterospecific song in female house sparrows: an in vivo positron emission tomography study. PLoS One 12(8):e0182875
LeBlanc MM, Goode CT, MacDougall-Shackleton EA, Maney DL (2007) Estradiol modulates brainstem catecholaminergic cell groups and projections to the auditory forebrain in a female songbird. Brain Res 1171:93–103
Lee V, Pawlisch BA, Macedo-Lima M, Remage-Healey L (2018) Norepinephrine enhances song responsiveness and encoding in the auditory forebrain of male zebra finches. J Neurophysiol 119(1):209–220
London SE, Monks DA, Wade J, Schlinger BA (2006) Widespread capacity for steroid synthesis in the avian brain and song system. Endocrinol 147(12):5975–5987
Lucas JR, Freeberg TM, Krishnan A, Long GR (2002) A comparative study of avian auditory brainstem responses: correlations with phylogeny and vocal complexity, and seasonal effects. J Comp Physiol A Neuroethol Sens Neural Behav Physiol 188(11–12):981–992
Lynch KS, Wilczynski W (2008) Reproductive hormones modify reception of species-typical communication signals in a female anuran. Brain Behav Evol 71(2):143–150
Lynch KS, Diekamp B, Ball GF (2012) Colocalization of immediate early genes in catecholamine cells after song exposure in female zebra finches (Taeniopygia guttata). Brain Behav Evol 79(4):252–260
Maney DL, Rodriguez-Saltos CA (2016) Hormones and the incentive salience of birdsong. In: Bass AH (ed) Hearing and hormones, vol 57. Springer, Heidelberg, pp 101–132
Maney DL, Richardson RD, Wingfield JC (1997) Central administration of chicken gonadotropin-releasing hormone-II enhances courtship behavior in a female sparrow. Horm Behav 32(1):11–18
Maney DL, Cho E, Goode CT (2006) Estrogen-dependent selectivity of genomic responses to birdsong. Eur J Neurosci 23(6):1523–1529
Mangiamele LA, Gomez JR, Curtis NJ, Thompson RR (2017) GPER/GPR30, a membrane estrogen receptor, is expressed in the brain and retina of a social fish (Carassius auratus) and colocalizes with isotocin. J Comp Neurol 525(2):252–270
Marler P (2008) Birdsong and monkey talk: an ethological journey. In: Zeigler HP, Marler P (eds) Neuroscience of birdsong. Cambridge University Press, New York, pp 449–462
Marler P, Peters S, Wingfield J (1987) Correlations between song acquisition, song production, and plasma-levels of testosterone and estradiol in sparrows. J Neurobiol 18(6):531–548
Marler P, Peters S, Ball GF, Dufty AM, Wingfield JC (1988) The role of sex steroids in the acquisition and production of birdsong. Nature 336(6201):770–772
Maruska, K. P., Becker, L., Neboori, A., & Fernald, R. D. (2013). Social descent with territory loss causes rapid behavioral, endocrine and transcriptional changes in the brain. J Exp Biol, 216(Pt 19), 3656–3666
Matragrano LL, Sanford SE, Salvante KG, Beaulieu M, Sockman KW, Maney DL (2012a) Estradiol-dependent modulation of serotonergic markers in auditory areas of a seasonally breeding songbird. Behav Neurosci 126(1):110–122
Matragrano LL, Beaulieu M, Phillip JO, Rae AI, Sanford SE, Sockman KW, Maney DL (2012b) Rapid effects of hearing song on catecholaminergic activity in the songbird auditory pathway. PLoS One 7(6):e39388
Meitzen J, Perkel DJ, Brenowitz EA (2007a) Seasonal changes in intrinsic electrophysiological activity of song control neurons in wild song sparrows. J Comp Physiol A Neuroethol Sens Neural Behav Physiol 193(6):677–683
Meitzen J, Moore IT, Lent K, Brenowitz EA, Perkel DJ (2007b) Steroid hormones act transsynaptically within the forebrain to regulate neuronal phenotype and song stereotypy. J Neurosci 27(44):12045–12057
Mello CV, Vicario DS, Clayton DF (1992) Song presentation induces gene expression in the songbird forebrain. Proc Natl Acad Sci U S A 89(15):6818–6822
Metzdorf R, Gahr M, Fusani L (1999) Distribution of aromatase, estrogen receptor, and androgen receptor mRNA in the forebrain of songbirds and nonsongbirds. J Comp Neurol 407(1):115–129
Naftolin, F., Ryan, K. J., Davies, I. J., Reddy, V. V., Flores, F., Petro, Z., Kuhn, M., White, R. J., Takaoka, Y., & Wolin, L. (1975). The formation of estrogens by central neuroendocrine tissues. Recent Progress in Hormone Research, Proceedings of the 1996 Conference, Vol 52, 31, 295–319
Noirot IC, Adler HJ, Cornil CA, Harada N, Dooling RJ, Balthazart J, Ball GF (2009) Presence of aromatase and estrogen receptor alpha in the inner ear of zebra finches. Hear Res 252(1–2):49–55
Nottebohm F, Arnold A (1976) Sexual dimorphism in vocal control areas of the songbird brain. Science 194(4261):211–213
Nottebohm F, Nottebohm ME, Crane LA, Wingfield JC (1987) Seasonal-changes in gonadal hormone levels of adult male canaries and their relation to song. Behav Neural Biol 47(2):197–211
Okuyama T, Suehiro Y, Imada H, Shimada A, Naruse K, Takeda H, Kubo T, Takeuchi H (2011) Induction of c-fos transcription in the medaka brain (Oryzias latipes) in response to mating stimuli. Biochem Biophys Res Commun 404(1):453–457
Orchinik M, Murray TF, Moore FL (1991) A corticosteroid receptor in neuronal membranes. Science 252(5014):1848–1851
Pawlisch BA, Remage-Healey L (2015) Neuroestrogen signaling in the songbird auditory cortex propagates into a sensorimotor network via an ‘interface’ nucleus. Neuroscience 284:522–535
Pawlisch BA, Riters LV (2010) Selective behavioral responses to male song are affected by the dopamine agonist GBR-12909 in female european starlings (Sturnus vulgaris). Brain Res 1353:113–124
Peterson RS, Yarram L, Schlinger BA, Saldanha CJ (2005) Aromatase is pre-synaptic and sexually dimorphic in the adult zebra finch brain. Proc Roy Soc B-Biol Sci 272(1576):2089–2096
Petrulis A (2013) Chemosignals, hormones and mammalian reproduction. Horm Behav 63(5):723–741
Phillmore LS, Veysey AS, Roach SP (2011) Zenk expression in auditory regions changes with breeding condition in male black-capped chickadees (Poecile atricapillus). Behav Brain Res 225(2):464–472
Reddy VV, Naftolin F, Ryan KJ (1973) Aromatization in the central nervous system of rabbits: effects of castration and hormone treatment. Endocrinology 92(2):589–594
Remage-Healey L (2014a) Frank Beach award winner: steroids as neuromodulators of brain circuits and behavior. Horm Behav 66(3):552–560
Remage-Healey, L. (2014b)
Remage-Healey L, Joshi NR (2012) Changing neuroestrogens within the auditory forebrain rapidly transform stimulus selectivity in a downstream sensorimotor nucleus. J Neurosci 32(24):8231–8241
Remage-Healey L, Maidment NT, Schlinger BA (2008) Forebrain steroid levels fluctuate rapidly during social interactions. Nat Neurosci 11(11):1327–1334
Remage-Healey L, Coleman MJ, Oyama RK, Schlinger BA (2010) Brain estrogens rapidly strengthen auditory encoding and guide song preference in a songbird. Proc Natl Acad Sci U S A 107(8):3852–3857
Remage-Healey L, Dong SM, Chao A, Schlinger BA (2012) Sex-specific, rapid neuroestrogen fluctuations and neurophysiological actions in the songbird auditory forebrain. J Neurophysiol 107(6):1621–1631
Remage-Healey L, Jeon SD, Joshi NR (2013) Recent evidence for rapid synthesis and action of oestrogens during auditory processing in a songbird. J Neuroendocrinol 25(11):1024–1031
Riebel K, Smallegange IM, Terpstra NJ, Bolhuis JJ (2002) Sexual equality in zebra finch song preference: evidence for a dissociation between song recognition and production learning. Proc Roy Soc B-Biol Sci 269(1492):729–733
Sakata, J. T., & Vehrencamp, S. L. (2012). Integrating perspectives on vocal performance and consistency. J Exp Biol, 215(Pt 2), 201–209
Saldanha CJ, Remage-Healey L, Schlinger BA (2013) Neuroanatomical distribution of aromatase in birds: cellular and subcellular analyses. In: Balthazart GBJ (ed) Brain aromatase, estrogens and behavior. Oxford, UK, Oxford, pp 100–114
Sanford SE, Lange HS, Maney DL (2010) Topography of estradiol-modulated genomic responses in the songbird auditory forebrain. Dev Neurobiol 70(2):73–86
Schlinger BA, Arnold AP (1992) Circulating estrogens in a male songbird originate in the brain. Proc Natl Acad Sci U S A 89(16):7650–7653
Schlinger B, Brenowitz EA (2008) Neural and hormonal control of birdsong. In: Pfaff DW (ed) Hormones, Brain and Behavior, vol 2. Elsevier, pp 897–941
Schlinger BA, Remage-Healey L (2012) Neurosteroidogenesis: insights from studies of songbirds. J Neuroendocrinol 24(1):16–21
Seredynski AL, Balthazart J, Ball GF, Cornil CA (2015) Estrogen receptor beta activation rapidly modulates male sexual motivation through the transactivation of metabotropic glutamate receptor 1a. J Neurosci 35(38):13110–13123
Sisneros JA, Bass AH (2003) Seasonal plasticity of peripheral auditory frequency sensitivity. J Neurosci 23(3):1049–1058
Sisneros JA, Forlano PM, Deitcher DL, Bass AH (2004) Steroid-dependent auditory plasticity leads to adaptive coupling of sender and receiver. Science 305(5682):404–407
Sizemore M, Perkel DJ (2008) Noradrenergic and GABAB receptor activation differentially modulate inputs to the premotor nucleus RA in zebra finches. J Neurophysiol 100(1):8–18
Smith GT, Brenowitz EA, Beecher MD, Wingfield JC (1997) Seasonal changes in testosterone, neural attributes of song control nuclei, and song structure in wild songbirds. J Neurosci 17(15):6001–6010
Soma KK, Schlinger BA, Wingfield JC, Saldanha CJ (2003) Brain aromatase, 5 alpha-reductase, and 5 beta-reductase change seasonally in wild male song sparrows: relationship to aggressive and sexual behavior. J Neurobiol 56(3):209–221
Soma KK, Bindra RK, Gee J, Wingfield JC, Schlinger BA (1999) Androgen-metabolizing enzymes show region-specific changes across the breeding season in the brain of a wild songbird. J Neurobiol 41(2):176–188
Srivastava DP, Evans PD (2013) G-protein oestrogen receptor 1: trials and tribulations of a membrane oestrogen receptor. J Neuroendocrinol 25(11):1219–1230
Tachikawa KS, Yoshihara Y, Kuroda KO (2013) Behavioral transition from attack to parenting in male mice: a crucial role of the vomeronasal system. J Neurosci 33(12):5120–5126
Tchernichovski O, Schwabl H, Nottebohm F (1998) Context determines the sex appeal of male zebra finch song. Anim Behav 55:1003–1010
Tinbergen N (1951) The study of instinct. Clarendon Press, Oxford Eng
Vahaba DM, Remage-Healey L (2015) Brain estrogen production and the encoding of recent experience. Curr Op Behav Sci 6:148–153
Vahaba DM, Remage-Healey L (2018) Neuroestrogens rapidly shape auditory circuits to support communication learning and perception: Evidence from songbirds. Horm Behav 104:77–87. https://doi.org/10.1016/j.yhbeh.2018.03.007
Vahaba DM, Macedo-Lima M, Remage-Healey L (2017) Sensory coding and sensitivity to local estrogens shift during critical period milestones in the auditory cortex of male songbirds. eNeuro 4(6)
Vasudevan N, Pfaff DW (2008) Non-genomic actions of estrogens and their interaction with genomic actions in the brain. Front Neuroendocrinol 29(2):238–257
Velho TA, Lu K, Ribeiro S, Pinaud R, Vicario D, Mello CV (2012) Noradrenergic control of gene expression and long-term neuronal adaptation evoked by learned vocalizations in songbirds. PLoS One 7(5):e36276
Vyas A, Harding C, McGowan J, Snare R, Bogdan D (2008) Noradrenergic neurotoxin, N-(2-chloroethyl)-N-ethyl-2-bromobenzylamine hydrochloride (DSP-4), treatment eliminates estrogenic effects on song responsiveness in female zebra finches (Taeniopygia guttata). Behav Neurosci 122(5):1148–1157
Wingfield JC, Farner DS (1976) Plasma Lh and sex steroids in whitecrowned sparrow, Zonotrichia-Leucophrys-Pugetensis. Amer Zool 16(2):257–257
Wingfield JC, Ball GF, Dufty AM, Hegner RE, Ramenofsky M (1987) Testosterone and aggression in birds. Amer Sci 75(6):602–608
Woolley SC, Doupe AJ (2008) Social context-induced song variation affects female behavior and gene expression. PLoS Biol 6(3):e62
Zigmond RE, Notteboh F, Pfaff DW (1973) Androgen-concentrating cells in midbrain of a songbird. Science 179(4077):1005–1007
Acknowledgements
Preparation of this chapter was supported in part by NIH R01NS082179 and NSF IOS 1354906.
Compliance with Ethics Requirements
Luke Remage-Healey declares that he has no conflict of interest.
Author information
Authors and Affiliations
Corresponding author
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 2020 Springer Nature Switzerland AG
About this chapter
Cite this chapter
Remage-Healey, L. (2020). Hormonal Regulation of Avian Auditory Processing. In: Sakata, J., Woolley, S., Fay, R., Popper, A. (eds) The Neuroethology of Birdsong. Springer Handbook of Auditory Research, vol 71. Springer, Cham. https://doi.org/10.1007/978-3-030-34683-6_6
Download citation
DOI: https://doi.org/10.1007/978-3-030-34683-6_6
Published:
Publisher Name: Springer, Cham
Print ISBN: 978-3-030-34682-9
Online ISBN: 978-3-030-34683-6
eBook Packages: Biomedical and Life SciencesBiomedical and Life Sciences (R0)