Publications

Peer-Reviewed Publications

Joyce, W. (2026). The Molecular Foundations of High Heart Rates: Sarcomeric Protein Adaptations. Physiology. In press. https://doi.org/10.1152/physiol.00021.2025

Joyce, W., He, K., Zhang, M., Ogunsola, S., Wu, X., Joseph, K. T., Bogomolny, D., Yu, W., Springer, M. S., Xie, J., Signore, A. V., & Campbell, K. L. (2024). Genetic excision of the regulatory cardiac troponin I extension in high–heart rate mammal clades. Science, 385(6716), 1466–1471. https://doi.org/10.1126/science.adi8146

Joyce, W., Shiels, H. A., & Franklin, C. E. (2024). The integrative biology of the heart: Mechanisms enabling cardiac plasticity. Journal of Experimental Biology, 227(20), jeb249348. https://doi.org/10.1242/jeb.249348

Joyce, W. (2023). Evolutionary loss of the β1-adrenergic receptor in salmonids. General and Comparative Endocrinology, 338, 114279. https://doi.org/10.1016/j.ygcen.2023.114279

Joyce, W. (2023). Muscle growth and plasticity in teleost fish: The significance of evolutionarily diverse sarcomeric proteins. Reviews in Fish Biology and Fisheries, 33(4), 1311–1327. https://doi.org/10.1007/s11160-023-09800-8

Joyce, W., Ripley, D. M., Gillis, T., Black, A. C., Shiels, H. A., & Hoffmann, F. G. (2023). A Revised Perspective on the Evolution of Troponin I and Troponin T Gene Families in Vertebrates. Genome Biology and Evolution, 15(1), evac173. https://doi.org/10.1093/gbe/evac173

Joyce, W., Warwicker, J., Shiels, H. A., & Perry, S. F. (2023). Evolution and divergence of teleost adrenergic receptors: Why sometimes ‘the drugs don’t work’ in fish. Journal of Experimental Biology, 226(19), jeb245859. https://doi.org/10.1242/jeb.245859

Boukens, B. J. D., Joyce, W., Kristensen, D. L., Hooijkaas, I., Jongejan, A., Wang, T., & Jensen, B. (2022). Catecholamines are key modulators of ventricular repolarization patterns in the ball python (Python regius). The Journal of General Physiology, 154(2), e202012761. https://doi.org/10.1085/jgp.202012761

Costello, L. M., García-Párraga, D., Crespo-Picazo, J. L., Codd, J. R., Shiels, H. A., & Joyce, W. (2022). Absence of atrial smooth muscle in the heart of the loggerhead sea turtle (Caretta caretta): A re-evaluation of its role in diving physiology. Journal of Experimental Biology, jeb.244864. https://doi.org/10.1242/jeb.244864

Joyce, W., Pan, Y. K., Garvey, K., Saxena, V., & Perry, S. F. (2022). Regulation of heart rate following genetic deletion of the ß1 adrenergic receptor in larval zebrafish. Acta Physiologica, 235(4), e13849. https://doi.org/10.1111/apha.13849

Joyce, W., & Wang, T. (2022). Regulation of heart rate in vertebrates during hypoxia: A comparative overview. Acta Physiologica, 234(3). https://doi.org/10.1111/apha.13779

Kunert, E., Joyce, W., Pan, Y. K., Chen, A., Perry, S. F., & Gilmour, K. M. (2022). The role of cytosolic carbonic anhydrase Ca17a in cardiorespiratory responses to CO2 in developing zebrafish (Danio rerio). American Journal of Physiology-Regulatory, Integrative and Comparative Physiology. https://doi.org/10.1152/ajpregu.00050.2022

Smith, B., Crossley, D. A., Wang, T., & Joyce, W. (2022). No evidence for pericardial restraint in the snapping turtle (Chelydra serpentina) following pharmacologically induced bradycardia at rest or during exercise. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 322(5), R389–R399. https://doi.org/10.1152/ajpregu.00004.2022

Bard, B., Dodge, A., Joyce, W., Lawrence, M., Cooke, S. J., & Gilmour, K. M. (2021). Elevated cortisol lowers thermal tolerance but results in limited cardiac remodelling in rainbow trout (Oncorhynchus mykiss) experiencing chronic social stress. Journal of Experimental Biology, jeb.238683. https://doi.org/10.1242/jeb.238683

Joyce, W., & Axelsson, M. (2021). Regulation of splenic contraction persists as a vestigial trait in white-blooded Antarctic fishes. Journal of Fish Biology, 98(1), 287–291. https://doi.org/10.1111/jfb.14579

Joyce, W., & Perry, S. F. (2021). Hif‐1α is not required for the development of cardiac adrenergic control in zebrafish ( Danio rerio ). Journal of Experimental Zoology Part A: Ecological and Integrative Physiology, 335(7), 623–631. https://doi.org/10.1002/jez.2507

Joyce, W., Scholman, K. T., Jensen, B., Wang, T., & Boukens, B. J. (2021). Α1-adrenergic stimulation increases ventricular action potential duration in the intact mouse heart. FACETS, 6, 823–836. https://doi.org/10.1139/facets-2020-0081

Joyce, W., & Wang, T. (2021). How cardiac output is regulated: August Krogh’s proto-Guytonian understanding of the importance of venous return. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 253, 110861. https://doi.org/10.1016/j.cbpa.2020.110861

Mandic, M., Joyce, W., & Perry, S. F. (2021). The evolutionary and physiological significance of the Hif pathway in teleost fishes. Journal of Experimental Biology, 224(18). https://doi.org/10.1242/jeb.231936

Nørgaard, S., Joyce, W., Jensen, M. F., Enok, S., Skovgaard, N., & Wang, T. (2021). Histamine exerts both direct H2-mediated and indirect catecholaminergic effects on heart rate in pythons. Journal of Comparative Physiology B. https://doi.org/10.1007/s00360-020-01338-6

O’Brien, K. M., Joyce, W., Crockett, E. L., Axelsson, M., Egginton, S., & Farrell, A. P. (2021). Resilience of cardiac performance in Antarctic notothenioid fishes in a warming climate. Journal of Experimental Biology, 224(10). https://doi.org/10.1242/jeb.220129

Jensen, B., Joyce, W., Gregorovicova, M., Sedmera, D., Wang, T., & Christoffels, V. M. (2020). Low incidence of atrial septal defects in nonmammalian vertebrates. Evolution & Development, 22(3), 241–256. https://doi.org/10.1111/ede.12322

Joyce, W., Crossley, D. A., Wang, T., & Jensen, B. (2020). Smooth Muscle in Cardiac Chambers is Common in Turtles and Extensive in the Emydid Turtle, Trachemys scripta. The Anatomical Record, 303(5), 1327–1336. https://doi.org/10.1002/ar.24257

Joyce, W., & Perry, S. F. (2020). Hypoxia inducible factor-1α knockout does not impair acute thermal tolerance or heat hardening in zebrafish. Biology Letters, 16(7), 20200292. https://doi.org/10.1098/rsbl.2020.0292

Joyce, W., & Wang, T. (2020). What determines systemic blood flow in vertebrates? The Journal of Experimental Biology, 223(4), jeb215335. https://doi.org/10.1242/jeb.215335

Boukens, B. J. D., Kristensen, D. L., Filogonio, R., Carreira, L. B. T., Sartori, M. R., Abe, A. S., Currie, S., Joyce, W., Conner, J., Opthof, T., Crossley, D. A., Wang, T., & Jensen, B. (2019). The electrocardiogram of vertebrates: Evolutionary changes from ectothermy to endothermy. Progress in Biophysics and Molecular Biology, 144, 16–29. https://doi.org/10.1016/j.pbiomolbio.2018.08.005

Joyce, W., Axelsson, M., & Wang, T. (2019). Contraction of atrial smooth muscle reduces cardiac output in perfused turtle hearts. The Journal of Experimental Biology, 222(Pt 6), jeb199828. https://doi.org/10.1242/jeb.199828

Joyce, W., Egginton, S., Farrell, A. P., & Axelsson, M. (2019). Adrenergic and adenosinergic regulation of the cardiovascular system in an Antarctic icefish: Insight into central and peripheral determinants of cardiac output. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 230, 28–38. https://doi.org/10.1016/j.cbpa.2018.12.012

Joyce, W., White, D. W., Raven, P. B., & Wang, T. (2019). Weighing the evidence for using vascular conductance, not resistance, in comparative cardiovascular physiology. Journal of Experimental Biology, 222(6), jeb197426. https://doi.org/10.1242/jeb.197426

Joyce, W., Williams, C. J. A., Iversen, S., Henriksen, P. G., Bayley, M., & Wang, T. (2019). The effects of endogenous and exogenous catecholamines on hypoxic cardiac performance in red-bellied piranhas. Journal of Experimental Zoology. Part A, Ecological and Integrative Physiology, 331(1), 27–37. https://doi.org/10.1002/jez.2233

Wang, T., Joyce, W., & Hicks, J. W. (2019). Similitude in the cardiorespiratory responses to exercise across vertebrates. Current Opinion in Physiology, 10, 137–145. https://doi.org/10.1016/j.cophys.2019.05.007

Bundgaard, A., James, A. M., Joyce, W., Murphy, M. P., & Fago, A. (2018). Suppression of reactive oxygen species generation in heart mitochondria from anoxic turtles: The role of complex I S-nitrosation. Journal of Experimental Biology, jeb.174391. https://doi.org/10.1242/jeb.174391

Joyce, W., Axelsson, M., Egginton, S., Farrell, A. P., Crockett, E. L., & O’Brien, K. M. (2018). The effects of thermal acclimation on cardio-respiratory performance in an Antarctic fish (Notothenia coriiceps). Conservation Physiology, 6(1). https://doi.org/10.1093/conphys/coy069

Joyce, W., Crossley, J., Elsey, R. M., Wang, T., & Crossley, D. A. (2018). Contribution of active atrial contraction to cardiac output in anesthetized American alligators (Alligator mississippiensis). Journal of Experimental Biology, jeb.178194. https://doi.org/10.1242/jeb.178194

Joyce, W., Egginton, S., Farrell, A. P., Crockett, E. L., O’Brien, K. M., & Axelsson, M. (2018). Exploring nature’s natural knockouts: In vivo cardiorespiratory performance of Antarctic fishes during acute warming. Journal of Experimental Biology, jeb.183160. https://doi.org/10.1242/jeb.183160

Joyce, W., Elsey, R. M., Wang, T., & Crossley, D. A. (2018). Maximum heart rate does not limit cardiac output at rest or during exercise in the American alligator (Alligator mississippiensis). American Journal of Physiology. Regulatory, Integrative and Comparative Physiology, 315(2), R296–R302. https://doi.org/10.1152/ajpregu.00027.2018

Joyce, W., Miller, T. E., Elsey, R. M., Wang, T., & Crossley, D. A. (2018). The effects of embryonic hypoxic programming on cardiovascular function and autonomic regulation in the American alligator (Alligator mississippiensis) at rest and during swimming. Journal of Comparative Physiology B, 188(6), 967–976. https://doi.org/10.1007/s00360-018-1181-2

Joyce, W., Williams, C. J. A., Crossley, D. A., & Wang, T. (2018). Venous pressures and cardiac filling in turtles during apnoea and intermittent ventilation. Journal of Comparative Physiology. B, Biochemical, Systemic, and Environmental Physiology, 188(3), 481–490. https://doi.org/10.1007/s00360-017-1132-3

Filogonio, R., Joyce, W., & Wang, T. (2017). Nitrergic cardiovascular regulation in the African lungfish, Protopterus aethiopicus. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 207, 52–56. https://doi.org/10.1016/j.cbpa.2016.12.030

Joyce, W., Axelsson, M., & Wang, T. (2017). Autoregulation of cardiac output is overcome by adrenergic stimulation in the anaconda heart. The Journal of Experimental Biology, 220(3), 336–340. https://doi.org/10.1242/jeb.149237

Joyce, W., Axelsson, M., Altimiras, J., & Wang, T. (2016). In situ cardiac perfusion reveals interspecific variation of intraventricular flow separation in reptiles. Journal of Experimental Biology, 219(14), 2220–2227. https://doi.org/10.1242/jeb.139543

Joyce, W., Ozolina, K., Mauduit, F., Ollivier, H., Claireaux, G., & Shiels, H. A. (2016). Individual variation in whole-animal hypoxia tolerance is associated with cardiac hypoxia tolerance in a marine teleost. Biology Letters, 12(1), 20150708. https://doi.org/10.1098/rsbl.2015.0708

Joyce, W., Simonsen, M., Gesser, H., & Wang, T. (2016). The effects of hypoxic bradycardia and extracellular HCO3-/CO2 on hypoxic performance in the eel heart. Journal of Experimental Biology, 219(3), 302–305. https://doi.org/10.1242/jeb.130971

Joyce, W., Gesser, H., Bayley, M., & Wang, T. (2015). Anoxia and Acidosis Tolerance of the Heart in an Air-Breathing Fish (Pangasianodon hypophthalmus). Physiological and Biochemical Zoology, 88(6), 648–659. https://doi.org/10.1086/682701

Joyce, W., Gesser, H., & Wang, T. (2014). Purinoceptors exert negative inotropic effects on the heart in all major groups of reptiles. Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology, 171, 16–22. https://doi.org/10.1016/j.cbpa.2014.02.005

Joyce, W., & Wang, T. (2014). Adenosinergic regulation of the cardiovascular system in the red-eared slider Trachemys scripta. Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology, 174, 18–22. https://doi.org/10.1016/j.cbpa.2014.04.002

Book Chapters

Jensen, B., Joyce, W., Wang, T., Ryerson, B., & Whitenack, L. B. (2025). Circulatory System. Comparative Vertebrate and Human Anatomy: Ecology, Evolution, and Function. https://pressbooks.palni.org/comparativevertebrateandhumananatomy/chapter/circulatory-system/

Joyce, W., & Brijs, J. (2024). Integrated responses of the circulatory system to digestion. In S. L. Alderman & T. E. Gillis (Eds.), Encyclopedia of Fish Physiology (Second Edition) (pp. 302–311). Academic Press. https://doi.org/10.1016/B978-0-323-90801-6.00124-5

Joyce, W., & Brijs, J. (2024). The gastrointestinal circulation. In S. L. Alderman & T. E. Gillis (Eds.), Encyclopedia of Fish Physiology (Second Edition) (pp. 147–157). Academic Press. https://doi.org/10.1016/B978-0-323-90801-6.00123-3

Wang, T., Filogonio, R., & Joyce, W. (2023). Evolution of the cardiovascular autonomic nervous system in vertebrates. In I. Biaggioni, K. Browning, G. Fink, J. Jordan, P. A. Low, & J. F. R. Paton (Eds.), Primer on the Autonomic Nervous System (4th ed., pp. 03–09). Elsevier Inc./Academic Press.

Editorials, Correspondences and Outreach

Borowiec, B. G., Joyce, W., & Glover, C. N. (2025). Editorial: Methods in Comparative Biochemistry and Physiology - Challenges and Opportunities. Comparative Biochemistry and Physiology. Part B, Biochemistry & Molecular Biology, 278, 111094. https://doi.org/10.1016/j.cbpb.2025.111094

Glover, C. N., Borowiec, B. G., & Joyce, W. (2024). Editorial: Common methodological issues in comparative biochemistry and physiology. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 111697. https://doi.org/10.1016/j.cbpa.2024.111697

Joyce, W. (2023) ECR Spotlight – William Joyce. J. Exp. Biol. 226. jeb246776

Joyce, W. and Wang, T. (2020) Response to ‘Flow versus pressure?’ J. Exp. Biol. 223 jeb.229054

Joyce, W. and Wang, T. (2020) Response to ‘What makes the blood go around?’ J. Exp. Biol. 223 jeb.225003

Joyce, W. (2019). Why a sea snake needs a hole in its head. J. Exp. Biol. 222, jeb.193185.

Joyce, W. (2019). Andean birds with slow metabolism live longer. J. Exp. Biol. 222, jeb193011.

Joyce, W. (2019). Parenting is hot work for blue tits. J. Exp. Biol. 222, jeb192708.

Joyce, W. (2019). Kicking kangaroo rats deter rattlesnakes. J. Exp. Biol. 222, jeb192831.

Joyce, W. (2018). A breath of fresh air for lungless salamanders. J. Exp. Biol. 221. jeb170308.

Joyce, W. (2018). Pythons sacrifice muscle to provide water for their eggs. J. Exp. Biol. 221: jeb170175.

Joyce, W. (2018). Insulin resistance cavefish avoid diabetes. J. Exp. Biol. 221: jeb170050.

Joyce, W., (2018). Flying is not for the small hearted. J. Exp. Biol. 221:  jeb169912.

Joyce, W., (2017). White-nose syndrome dehydrates bats. J. Exp. Biol. 220, 4326–4326. 

Joyce, W., (2017). More haste, less speed for quolls. J. Exp. Biol. 220, 3003–3004.

Joyce, W. (2017). Fidgety embryos grow longer limbs. J. Exp. Biol. 220, 1936.

Joyce, W. (2017). An ancient origin for the diaphragm. J. Exp. Biol. 220, 737. 

Joyce, W. (2016). The shark heart crowned. J. Exp. Biol. 219, 3674-3675. 

Joyce, W. (2016). From pinnipeds to people: divers have elastic arteries. J. Exp. Biol. 219, 2579. 

Joyce, W. (2016). Taurine tunes cuttlefish cardiac output. J. Exp. Biol. 219, 1588-1588.

Joyce, W. (2016). Anaemic starlings have fewer chicks (sometimes). J. Exp. Biol. 219, 614–614.