Announcing the 2025 Journal of Experimental Biology Outstanding Paper Prize shortlist and winner

If you're a fan of comparative physiology, clicking on a new issue of Journal of Experimental Biology can feel like being let loose in an opulent store of treats. From stories about surfing zooplankton and caterpillar sound effects to the map sense of turtles and the impact of climate change, you can never be sure what you'll get, but it will be extraordinary. ‘JEB's strength lies in embracing diversity, from the animals studied, questions asked and approaches taken’, says Craig Franklin, JEB Editor-in-Chief. Behind each remarkable piece of research lie the personal stories of the early-career scientists that nurtured animals and painstakingly recorded their behaviour and physiology. ‘Supporting early-career researchers is fundamental to JEB's identity and our investment in the future of comparative physiology and biomechanics’, says Franklin. Each year, JEB recognises the achievements of the journal's early-career researchers, who make immense contributions to the research published in the journal, through the JEB Outstanding Paper Prize. Launched in 2005 in memory of former JEB Editor-in-Chief Robert (Bob) G. Boutilier, the shortlist provides a glimpse of the diverse topics covered by the journal, introducing the early-career researchers behind each remarkable discovery. The 2025 shortlist is a treat of scientific delight, revealing the creativity and ingenuity of the early-career researchers that brought each study to completion (Box 1). Box 1. Shortlisted articles for the 2025 JEB Outstanding Paper PrizeN. D. Antonson, S. Ogunbiyi, M. Champigneulle, T. J. Roberts, F. Goller and M. J. Fuxjager: Neuromuscular coordination of movement and breathing forges a hammer-like mechanism for woodpecker drilling.O. E. Demuth, J. R. Hutchinson, V. La Barbera, S. E. Warner and D. J. Field: Soft tissue constraints on joint mobility in the avian shoulder.D. Giacometti and G. J. Tattersall: Behavioural evidence of a humidistat: a temperature-compensating mechanism of hydroregulation in spotted salamanders.J. L. Huisken and S. M. Rehan: Transcriptomic correlates of nutritional manipulation in a facultatively social bee.K. Q. Karkosiak, R. Z. Schwartz, H. King and T. A. Blackledge: Spider egg sacs reveal how pockets of air can be used to conserve water.S. D. Lee, S. Wang, D. Kuang, E. K. Wang, J. K. Yim, N. H. Hunt, R. S. Fearing, H. S. Stuart and R. J. Full: Free-ranging squirrels perform stable, above-branch landings by balancing using leg force and nonprehensile foot torque.A. M. Martin, D. Cordero-De La Cruz and L. Swierk: High dissolved oxygen extends dive duration and suggests physical gill use in a vertebrate.T. Mulder, Y. Yang, E. Morley, T. E. Miller, D. Hending, G. K. Taylor and B. Mortimer: Constraints on percussive seismic signals in a noisy environment by European fiddler crabs, Afruca tangeri.A. B. Thies, M. Rangarajan-Paul, D. Wangpraseurt and M. Tresguerres: Co-option of immune and digestive cellular machinery to support photosymbiosis in amoebocytes of the upside-down jellyfish Cassiopea xamachana.In the first of the shortlisted papers (Antonson et al., 2025), Nicholas Antonson (Brown University, USA) and colleagues from Brown University and The University of Utah, USA, investigated how downy woodpeckers (Dryobates pubescens) coordinate and control their muscles and breathing while pecking wood. ‘This project was something that I had long wanted to do’, says PI Matthew Fuxjager, who worked with Antonson during his postdoctoral fellowship. ‘Nick's quantitative skills are very impressive’, says Fuxjager, recalling that Antonson figured out how to visualise and analyse the complex data to test their ideas about how woodpeckers use their muscles during their forceful impacts. ‘This paper stood out for me for a few reasons’, says nominating Editor Katie Gilmour, adding that the experiments performed by Antonson – recording muscle electrical activity and air-sac pressure while the birds were actively moving and drilling wood – were extremely challenging.Switching focus from a unique activity performed by a few bird species to an almost ubiquitous manoeuvre performed by most – wing flapping – Oliver Demuth and Daniel Field (University of Cambridge, UK) with John Hutchinson and colleagues from the Royal Veterinary College (RVC), UK, investigated the range of motion of the partridge shoulder joint (Demuth et al., 2025). ‘The impetus for the study is the perennial challenge of drawing robust conclusions about joint range of motion for extinct vertebrate animals’, says Field, explaining that simulations of the movements of fossil joints, including the soft tissues that have been lost, are essential to understand the evolution of modern birds. Describing how Demuth delicately manoeuvred the partridges’ shoulder joints in an X-ray machine to follow the full range of motion, Hutchinson recalls how Demuth then painstakingly analysed the joint's movements in 3D with support from Vittorio La Barbera and Sharon Warner (both at RVC). ‘This study offers major strides in revealing the essential roles played by the soft tissue in a single joint's mobility using a cutting-edge integration of computational modelling and sophisticated experimental measurements’, says nominating Editor Matt McHenry.In the next shortlisted article, nominated by Craig Franklin, Danilo Giacometti and Glenn Tattersall from Brock University, Canada, investigated the ability of amphibians to regulate hydration (Giacometti and Tattersall, 2025). ‘This study stands out for revealing an unexpected sensory mechanism in that salamanders appear to detect rates of evaporative water loss itself, not just ambient humidity’, says Franklin. Reflecting on Giacometti's role in the study, Tattersall says, ‘Danilo was the primary drive for the research, writing and design’, adding that he was impressed by Giacometti's ‘willingness to pursue difficult, cross-disciplinary questions requiring mastery of biophysical, physiological and behavioural concepts’. Having overseen peer review of the research, Franklin is excited by the study because it, ‘transforms our view of amphibian hydro-regulation from passive tolerance to active, informed behavioural regulation’.Moving from the realm of amphibians to the insect world, the next article shortlisted for the JEB Outstanding Paper Prize explores the mechanisms underlying the extraordinarily cooperative social structure of eastern small carpenter bees (Ceratina calcarata). In their study, Jesse Huisken and Sandra Rehan investigated whether diet underlies the role differences between dwarf eldest daughters, which are small and destined to serve their younger sisters, and regular sisters that have the potential to go on and found future colonies (Huisken and Rehan, 2025). However, when the duo investigated the differences in brain gene activation between the two castes, they realised that social experience and environment have more of an impact on the firstborn's destiny than diet or birth order. ‘This study stood out because it goes beyond simple correlative analysis to do an experimental manipulation and then track impact at the transcriptomic level to understand the mechanisms underlying the observed changes in social roles’, says nominating Editor Trish Schulte.From bee youngsters to spider eggs, nurturing the next generation is a key JEB theme. In their shortlisted paper, Katherine Karkosiak and Todd Blackledge (The University of Akron, USA) with Ravi Schwartz and Hunter King (Rutgers University – Camden, USA) investigated the permeability of spider egg sac silk, to determine how much protection the silk offers eggs from dehydration (Karkosiak et al., 2025). ‘We show that even though egg sacs look solid to our eyes, water vapour easily moves across them through tiny holes’, says Blackledge, which was a surprise. The team eventually realised that the egg sacs protect the eggs within by slowing the diffusion of water across the gap between the eggs and silk case and by shielding the eggs from moving air. ‘What makes this article exceptional is the authors’ systematic approach to answering a biological question’, says Franklin. And Blackledge, the PI on the study, was impressed by Karkosiak's resilience. ‘Kate showed immense scientific grit by transforming initial “failure” into a key insight for how water vapour moves across porous surfaces’, he says, adding that Schwartz's development of the physical model that explains how water diffuses through porous structures was a fantastic example of ‘integrated teamwork at its best!’.In a remarkable study investigating the landings of free-ranging fox squirrels on the University of California, Berkeley campus (USA), students Sebastian Lee, Stanley Wang and Duyi (Tina) Kuang – at Berkeley at the time – trained wild squirrels to jump into a rig that could measure the forces exerted as they touched down, to learn how squirrels, which land palm down on branches, avoid slipping off during botched landings (Lee et al., 2025). When the squirrels misjudged a short landing and overshot, they extended their rear legs, increasing their inertia to brake and prevent themselves from toppling forward. In contrast, the squirrels that undershot when trying to land a leap that was too far pulled themselves into a faster moving tight ball to swing themselves back up onto the branch. PI Robert Full says, ‘The students’ efforts paid off by discovering the secrets of dynamic balance, leading to the first legged robot that could land upright on a branch’. And nominating Editor Monica Daley says that she liked the paper because, ‘it involved challenging and creative experiments that required ingenuity and patience to collect data from wild animals using custom instrumentation’.An extraordinary investigation by graduate student Alexandra Martin and Lindsey Swierk (Binghamton University, USA) and then-undergraduate Diane Cordero-De La Cruz (University of California San Marcos, USA) reveals that water anoles (Anolis aquaticus) may be the first terrestrial vertebrate to breathe oxygen from water using a bubble as a physical gill to extract the gas from water (Martin et al., 2025). Filming the anoles submerged in normal, poorly oxygenated and super-oxygenated water as the reptiles rebreathed oxygen from an air bubble clinging to their foreheads, the researchers obtained several pieces of evidence suggesting that the bubble could be absorbing oxygen from the surrounding water to supplement the anoles’ oxygen supply. ‘Allie led this work as part of her doctoral dissertation’, says Swierk, adding that Martin invested a great deal of persistence and creativity, while Cordero-De La Cruz showed exceptional promise as an undergraduate researcher. JEB Editor Almut Kelber, who oversaw peer review of the research, said, ‘I like their simple but convincing and well-controlled experiment, their tests in more realistic environments with fast or gently flowing water and their caution pointing out that more data may have to be collected’.Editor Sheila Patek's nomination for the 2025 JEB Outstanding Paper Prize shortlist investigated the messages communicated by male European fiddler crabs when they beat the sand with their claws and drop their bodies to pound the sand (Mulder et al., 2025). Tom Mulder, Ellen Morley and Thomas Miller (University of Oxford, UK) recorded over 8000 percussive crab serenades with geophones embedded in a Portuguese beach over 2 months, revealing that the crabs with the largest claws stood out from the crowd by producing the loudest thuds. In addition, the percussive beats were not tuned to a particular tone, which could be an advantage in the noisy environment of the seashore, where wind and waves could drown out certain tones. ‘Tom led most aspects of this paper from inception to write up. This was particularly impressive given this was his first piece of academic fieldwork’, says PI Beth Mortimer (University of Oxford). And Patek pays tribute to the work saying, ‘This type of acoustic signalling in a noisy environment is very challenging to study, especially to disentangle individual effects and the interaction between the sound-producing mechanism and the changing environmental conditions’.In the final shortlisted paper of 2025, Angus Thies, Maitri Rangarajan-Paul, Daniel Wangpraseurt and Martin Tresguerres report how the cells that house symbiotic algae in the bodies of upside-down jelly fish have independently evolved the same mechanism as corals, anemones and giant clams to enhance algal production of sugars and oxygen, which are provided to the jellyfish (Thies et al., 2025). ‘There are so many cool things about this study’, says nominating Editor Pat Wright, who oversaw peer review of the article, recalling that the team used a variety of experimental approaches to unpick details about the ion transport mechanisms that permit the jellyfish to use algae kidnapped from their diet to directly fuel their tissues. PI Tresguerres is particularly proud of graduate student Thies, who also mentored Rangarajan-Paul during the research. ‘Angus did an amazing job, including contributing to the original idea and leading all of the experiments’, says Tresguerres.After reviewing the nine nominated research articles, the Editors are delighted to announce that Nicholas Antonson, Brown University, USA, has been awarded the 2026 JEB Outstanding Paper Prize for his paper, ‘Neuromuscular coordination of movement and breathing forges a hammer-like mechanism for woodpecker drilling’ (Antonson et al., 2025). ‘The story is both compelling and novel, an impressive winner of the JEB Outstanding Paper Prize’, says Gilmour, who nominated the paper for the award. Reacting to the news, Antonson exclaimed, ‘I was elated!’, adding that the award was completely unexpected.‘The project was something Franz Goller and I cooked up several years ago’, says Fuxjager. ‘We had been having conversations about motor coordination of drilling and drumming, debating how complex the mechanisms likely were’, he adds, explaining that they were curious to know whether drilling was a whole-body affair and whether breathing was involved at all. Joining Fuxjager's lab in 2023 as a National Science Foundation postdoctoral fellow, Antonson set about collecting eight of the tiny woodpeckers for a 3-day stay in the lab. Then he filmed their manoeuvres and recorded their muscle activity as they pecked and drilled a piece of hardwood. ‘The most challenging aspect of this research was getting all the different types of data – electromyography, respiratory pressure and high-speed video – to line up correctly so we could reconstruct a complete picture of how the woodpeckers coordinate movement with breathing during pecking’, says Antonson. And Fuxjager recalls that Antonson's ability to simultaneously handle and visualise the complex datasets to unpick the fine detail was impressive. ‘I think the most exciting moment was when we noticed that the woodpeckers were exhaling through each strike they were producing’, says Antonson. Effectively, the birds were grunting – like tennis stars each time they strike a ball – which likely helps to boost the power of each blow. Even more impressively, the woodpeckers were able to grunt up to 13 times per second when drumming on wood to assert territory or attract a mate: ‘It's kind of like a musician keeping a crazy-fast beat: if you can do it perfectly, people notice. Woodpeckers do the same thing and the best drummers stand out amongst their rivals’, Antonson explains.‘It's really rewarding to see a project that has been stewing for a very long time get honoured as a significant contribution’, says Fuxjager, and Antonson has been excited by the wider reaction to the paper. ‘I even had to act out what the rapid “grunting” sounded like during an interview on [US National Public Radio's] Science Friday, which gave my friends and family a good chuckle’, he smiles. Looking to the future, Antonson is excited by the prospect of setting up his own lab and Fuxjager is optimistic for his success; ‘Nick is a very creative scientist who is full of new ideas’, he says.Reflecting on the full Outstanding Paper Prize shortlist, Franklin is excited by the biological diversity that it represents, showcasing groundbreaking research in vertebrates and invertebrates with implications for palaeontology, biomechanics, physiology and neuroethology. ‘This shortlist captures everything I love about JEB’, Franklin says with a smile. And he is impressed by the determination and commitment of all of the early-career researchers whose work has been highlighted. ‘With scientists of this calibre coming through, the future of comparative physiology and biomechanics is in extraordinarily capable hands’, says Franklin, looking toward another exciting year of inspiration and discovery with Journal of Experimental Biology.