Abstract
A recent rise in the global brewery sector has increased the demand for high-quality, late summer hops. The effects of ongoing and predicted climate change on the yield and aroma of hops, however, remain largely unknown. Here, we combine meteorological measurements and model projections to assess the climate sensitivity of the yield, alpha content and cone development of European hops between 1970 and 2050 CE, when temperature increases by 1.4 °C and precipitation decreases by 24 mm. Accounting for almost 90% of all hop-growing regions, our results from Germany, the Czech Republic and Slovenia show that hop ripening started approximately 20 days earlier, production declined by almost 0.2 t/ha/year, and the alpha content decreased by circa 0.6% when comparing data before and after 1994 CE. A predicted decline in hop yield and alpha content of 4–18% and 20–31% by 2050 CE, respectively, calls for immediate adaptation measures to stabilize an ever-growing global sector.
Similar content being viewed by others
Introduction
Beer is the world’s third most widely consumed beverage after water and tea1, and traditional beer brewing in central Europe dates back at least to the Neolithic period circa 3500–3100 BC2. In addition to water, malting barley and yeast, a much more expensive hop is needed to give beer its incomparable taste3. The specific hop aroma emerges from its bitter acid content and many other compounds, including essential oils and polyphenols4,5,6. Changes in alpha bitter acids affect the quality of hops7,8,9,10,11,12, and there has been a recent change in consumer preference towards beer aromas and flavors that heavily depend on high-quality hops13,14. Amplified by the ongoing craft beer popularity13, this trend contrasts with previous demands for lower alpha content14. The recent craft beer expansion therefore not only triggered new microbreweries but also boosted the demand for aromatic hops globally15,16. Although linkages between hop production and climate variation have been reported at local to regional scales9,10,17,18,19,20,21,22,23,24,25,26,27, relatively little is known about the possible, direct and indirect, effects of a predicted warmer and drier climate on the yield and alpha content of hops.
Since the cultivation of high-quality aroma hops is restricted to relatively small regions with suitable environmental conditions (Fig. 1), there is a serious risk that much of the production will be affected by individual heat waves or drought extremes that are likely to increase under global climate change28. Hop farmers can and have responded to climate change by relocating hop gardens to higher elevations and valley locations with higher water tables, building irrigation systems10, changing the orientation and spacing of crop rows, and even breeding more resistant varieties29. Changing the orientation of crop rows and combining irrigation with water-saving soil management practices have proven to be effective adaptation measures in viticulture30,31. It is important that the generative phase of hop plants occurs only in the appropriate photoperiod when sunshine duration is decreasing. This can be achieved by slowing plant growth via growth inhibitors or by building protective shading structures; which is, however, quite expensive. There is a similar problem in vineyards where shading by agrovoltaic panels has been introduced32,33. The higher probability of droughts can be partly mitigated by less frequent tillage and cultivation of hop fields, changes in fertilization and the use of row cover crops to support root growth34. A systematic and European-wide investigation of the impact of ongoing and predicted climate change on the quality and quantity of aroma hops is, however, still missing.
The top left map shows five sites representing 3 main German hop-growing regions (Spalt, Hallertau and Tettnang), the Zatec site in Czechia and Celje in Slovenia. The top right map indicates the approximate location of the main hop growing areas in Germany, Czechia and Slovenia as well as recognized hop-producing administrative regions in the remaining countries. The map also includes the locations of the 59 weather stations used in the analysis. The color of the dots responds to the hop acreage and the brown band to the optimal hop-growing zone.
Here, we show how temperature and precipitation control the yield, alpha content and cone development of aroma hops in Germany, the Czech Republic and Slovenia between 1970 and 2050 CE. We simulate the effect of weather conditions on hop yield and alpha content with a newly developed model. The model requires air temperature and precipitation records for input. Using simulations of future climate, we predict yield and alpha content. We then discuss how hop farmers can implement innovative adaptation measures to stabilize international markets under predicted global warming.
Results and discussion
Comparison of the average annual yield of European aroma hops during two independent periods, 1971–1994 and 1995–2018, reveals a significant production decrease in the range of 0.13–0.27 t/ha (Fig. 2). The average annual hop yield decreased after 1995 by 19.4% in Celje, 19.1% in Spalt, 13.7% in Hallertau, and 9.5% in Tettnang, whereas the production remained stable in Zatec (0.05 t/ha). In addition to the observed production decline, there were significant declines in the alpha content in all regions between from 0.46 and 1.86% (Fig. 2). The average alpha content decreased by 34.8% in Celje, 15.6% in Hallertau, 15% in Tettnang, 11.5% in Spalt, and 10.5% in Zatec. Hop yields between 1995 and 2018 exceeded the average yields from the 1971–1994 early period only once in Spalt and twice in Celje. Similarly, the average alpha content from the 1971–1994 early period was exceeded only one once in Celje and twice in Tettnang between 1995 and 2018. Declines in hop yields of more than 30% were recorded in 2000 and 2003. In 2006 and 2015, the decrease in alpha content was greater than 40%. Rising temperatures shifted the onset of the hop growing season by 13 days from 1970 to 2018. After 1995, the average onset of clone development (BBCH 71) occurred earlier compared to that in 1971–1994: 31 days earlier in Celje, 22 days earlier in Zatec, 16 days earlier in Hallertau and Spalt, and 13 days earlier in Tettnang (Fig. 2). These phenological changes shifted the critical ripening period towards the warmer part of the season, which had a negative impact on the alpha content.
To further assess the effects of changing weather conditions on the yields and alpha content of aromatic hops, we developed a parsimonious model simulating the variation in yields and alpha contents based on the difference in the precipitation and temperature from the optimal conditions during the growing season. High yields and alpha contents were obtained in years when weather conditions were close to optimum conditions, whereas low values occurred in years with extreme weather conditions. Hop yields increased with precipitation but decreased after ~15% of normal precipitation, whereas alpha contents decreased linearly with increasing temperatures. We found a statistically significant correlation (r = 0.41–0.73; p < 0.01) between the difference in the precipitation total from the optimum conditions in the growing season and hop yields (Supplementary Fig. 1). Moreover, a strong correlation was found between the average temperature at the time of heading and the alpha content (r = 0.61–0.78; p < 0.01; 1979–2018) (Supplementary Fig. 2). We found that sunshine duration at the time of heading was positively correlated with alpha content at p < 0.01. The lowest hop yields were negatively affected by a lack of precipitation, while the lowest values of alpha content were caused by extremely high temperatures. High temperatures and sunshine duration caused a sharp drop in alpha content in 2006 in all study areas. The estimated model risk of a decrease in alpha content was the consistently low production that occurred in 2006 in all hop regions in Europe. When temperature and light extremes occurred, most growing areas tended to be negatively affected, and production from other areas could not cover any losses.
Model projections for 2021–2050 suggested a decline in hop yields from 4.1–18.4% when compared to 1989–2018 (Fig. 3). A decrease of 20–30.8% was also projected for alpha content (Fig. 3). The most pronounced declines are expected to occur in the southern hop growing regions in southern Germany and Slovenia (Tettnang and Celje), while the more northern sites in Germany and the Czech Republic (Hallertau, Spalt and Zatec) are expected to experience less pronounced decreases in both parameters. The projected decrease will be caused mainly by rising temperatures and more frequent and severe droughts. These factors will most likely cause a significant decrease in alpha/ha production of 25.3–39.5% compared to current values (Fig. 3).
Changes in hop production and alpha content across Europe and the British Isles support our simulation results (Fig. 4). All scenarios predicted a decline in hop yields between 12% and 35% over 2021–2050 across all major hop growing regions in Europe, with Slovenia, Portugal, and Spain exhibiting the most pronounced declines (Supplementary Fig. 2). The alpha content was predicted to decrease considerably across all regions (Supplementary Fig. 2). Moderate decreases in yield and alpha content were predicted for Germany, the Czech Republic, and Poland, while the strongest declines in hop productivity were predicted for Portugal, Slovenia and Croatia (Supplementary Fig. 2).
Due to changes in climate and water availability for agriculture, some authors have addressed water stress and its influence on crops28,34,35,36,37. Quantification of the effect of deficit irrigation on hop yields, quality and profitability was performed in Washington State, USA. The results show that plants generally responded to water deficit with yield reductions. The 60% irrigation level caused reductions of 19–33% in the 2-year total yield, while the 80% irrigation level caused total 2-year yield changes from −14% to +2%. However, the quality of the hop cone was not affected. The concentrations of alpha-acids remained similar at all irrigation levels34.
In addition to the climatic factors described above, there are other external factors that can affect the yield and alpha content of aroma hops. These include the health condition of the hops, epigenetic adaptation and heredity of the hops, irrigation systems, harvest maturity, habitat conditions, and the regulation of hop growth by properly selected agricultural technology and fertilization24,38,39,40. The location of hop fields and suitable soil are very important, especially in the valleys of rivers and smaller streams, where the water table is generally more stable. The importance of drip irrigation combined with advanced irrigation planning technologies, such as the FORHOPS41 initiative for stabilizing hop yields has been successfully applied in the Zatec region, where irrigated hops have become dominant since 2015. Moreover, wetter and cooler locations have often been used for new hop fields, while open plateau fields have been reduced.
While assessing future climate and environmental impacts on the quality and quantity of aroma hops, the uncertainty associated with model simulations should also be noted. Increased CO2 concentrations could partially compensate for the effects of drought and support yield growth and leaf area index while improving water use efficiency42. However, this effect on hops is still under investigation, and we do not yet have enough evidence and knowledge. New findings in hop physiology, such as the beneficial effect of elevated CO2 on the primary metabolism of hop strobilus43 and the effects of vernalization and dormancy44 may help in the future to breed hops that are more resistant. Obtaining more detailed measurements and observations directly from hop gardens will allow more detailed seasonal climate analysis in the future and thus improve existing models.
Since agricultural droughts are projected to increase with high confidence in southern Europe and medium confidence in central Europe45, it will be necessary to expand the area of aroma hops by 20% compared to the current production area to compensate for a future decline in alpha content (and/or hop production). Some uncertainties also need to be considered, such as how hop growers will adopt climate-smart agriculture46 that seeks to increase sustainable productivity, strengthen farmer resilience, adapt marketing strategies to customers47, reduce agricultural greenhouse gas emissions and increase carbon sequestration. All factors will affect the economics and prospects for the future of aroma hop cultivation in Europe.
In summary, this study demonstrates a climate-induced decline in the quality and quantity of traditional aroma hops across Europe and calls for urgent adaptation measures to stabilize international market chains.
Methods
To identify the current and future impacts of climate change on aromatic hops, we selected important hop-growing areas in Germany (DE), Czechia (CZ) and Slovenia (SLO). These countries cover almost 90% of the total area of aromatic hop fields in Europe. The hop-farming regions of Hallertau (DE), Spalt (DE), Tettnang (DE), Zatec (in German, Saaz; CZ) and Celje (SLO) were selected for this study (Fig. 1). All study areas are located between 46–51°N and 9–15°E in regions with a mild continental climate. The average annual temperature varies between 8 and 10 °C, and the average annual precipitation fluctuates from 550 to 1050 mm. The altitudes of the hop fields vary between 200 and 500 m ASL, and the predominant soils are clay and loam. Hop acreages, average yields and alpha content (bitter substances in aromatic hops) for the individual areas were obtained from the Barth-Haas reports for 1970–201848. Daily precipitation totals, average daily temperatures, and onset of hop cone development were obtained from the German, Czech and Slovenian national meteorological services. Using regression kriging, which uses altitude as a predictor, we interpolated temperature and precipitation measurements to a resolution of 1*1 km. Daily sunshine duration totals were collected from EUMETSAT satellite observations SARAH 2.1 with a resolution of 0.05 * 0.05°49. The obtained layers were then used to calculate the average daily and monthly temperatures and precipitation totals for the selected hop-farming areas.
To assess the effects of weather conditions on the yields and alpha content of aroma hops, we developed a simple model (see SI) simulating the risk of lower yields based on the difference in the rainfall amounts from optimal conditions during the growing season and lower alpha contents based on the temperatures during cone development. Details of the models used are reported in the supplementary information. The formulation of the models is described here in Supplementary Table 1, and the results of the model calibration are shown in Supplementary Fig. 1.
Three global circulation models (GCMs) from the CMIP5 ensemble were used to represent the known variability in the rate of temperature change and precipitation patterns. The model CSIRO-MK36 (CSIRO) was used as the moderate estimate, GISS-E2-R-CC (GISS) represented a lower rate of temperature change and modest increase in dryness, and HadGEM2-ES (HadGEM) represented higher climate sensitivity with a hotter and drier climate. The representative concentration pathway, RCP4.5, was used for the construction of local-scale climate scenarios with the climate models. The climate projections from the GCMs were downscaled to local-scale daily weather by the LARS-WG 6.0 weather generator using the ELPIS dataset of site-specific parameters across Europe50. The combined average of the three global mean emissions models (CSIRO MK36; GISS-E2-R-CC; and HadGEM2-ES) was used to simulate the future evolution of yields and alpha contents in 2021–2050 (future) for comparison with those in 1989-2018 (present). The analysis is presented in detail for 5 selected sites and then for 59 ELPIS sites representing all EU and UK hop-producing regions.
Reporting summary
Further information on research design is available in the Nature Portfolio Reporting Summary linked to this article.
Data availability
All data that support the findings of this study are publicly available in FigShare (https://doi.org/10.6084/m9.figshare.23180342)51.
References
Nelson, M. The Barbarian’s Beverage: A History of Beer in Ancient Europe (Routledge, 2005).
McGovern, P. E. Uncorking the Past: The Quest for Wine, Beer, and Other Alcoholic Beverages (Univ. of California Press, 2009).
Lerro, M., Marotta, G. & Nazzaro, C. Measuring consumers’ preferences for craft beer attributes through best-worst scaling. Agric. Food Econ. 8, 1 (2020).
Roberts, T. R. Hops. In Brewing Materials and Processes (eds Bamforth, C. W.) 47–75 (Academic Press, 2016).
Goncalves, J., Figueira, J., Rodrigues, F. & Câmara, J. S. Headspace solid‐phase microextraction combined with mass spectrometry as a powerful analytical tool for profiling the terpenoid metabolomic pattern of hop‐essential oil derived from Saaz variety. J. Sep. Sci. 35, 2282–2296 (2012).
Astray, G., Gullón, P., Gullón, B., Munekata, P. E. S. & Lorenzo, J. M. Humulus lupulus L. as a natural source of functional biomolecules. Appl. Sci. 10, 5074 (2020).
Nance, M. R. & Setzer, W. N. Volatile components of aroma hops (Humulus lupulus L.) commonly used in beer brewing. J. Brew. Distill. 2, 16–22 (2011).
Forteschi, M. et al. Quality assessment of cascade hop (Humulus lupulus L.) grown in Sardinia. Eur. Food Res. Technol. 245, 863–871 (2019).
Lafontaine, S. R. & Shellhammer, T. H. How hoppy beer production has redefined hop quality and a discussion of agricultural and processing strategies to promote it. Tech. Q. 56, 1–12 (2019).
Mozny, M. et al. The impact of climate change on the yield and quality of Saaz hops in the Czech Republic. Agric. Meteorol. 149, 913–919 (2009).
Kučera, J. & Krofta, K. Mathematical model for prediction of alpha acid contents from meteorological data for’Saaz’aroma variety. ISHS Acta Hortic. 131–139 https://doi.org/10.17660/ActaHortic.2009.848.14 (2008).
Hieronymus, S. For the Love of Hops: The Practical Guide to Aroma, Bitterness, and the Culture of Hops. (Brewers Publications, a division of the Brewers Association, 2012).
Murray, D. W. & O’Neill, M. A. Craft beer: penetrating a niche market. Br. Food J. 114, 899–909 (2012).
Denby, C. M. et al. Industrial brewing yeast engineered for the production of primary flavor determinants in hopped beer. Nat. Commun. 9, 965–965 (2018).
Legun, K., Comi, M. & Vicol, M. New aesthetic regimes: the shifting global political ecology of aroma hops. Geoforum 128, 148–157 (2022).
Martin, A., Markhvida, M., Hallegatte, S. & Walsh, B. Socio-economic impacts of COVID-19 on household consumption and poverty. Econ. Disasters Clim. Change 4, 453–479 (2020).
Srečec, S., Ceh, B., Ciler, T. S. & Rus, A. F. Empiric mathematical model for predicting the content of alpha-acids in hop (Humulus lupulus L.) cv. Aurora. SpringerPlus 2, 59 (2013).
Srečec, S., Kvaternjak, I., Kaučić, D., Špoljar, A. & Erhatić, R. Influence of climatic conditions on accumulation of α-acids in hop clones. Agric. Conspec. Sci. 73, 161–166 (2008).
Wang, G. et al. Terpene biosynthesis in glandular trichomes of hop. Plant Physiol. 148, 1254–1266 (2008).
Barry, S., Muggah, E. M., McSweeney, M. B. & Walker, S. A preliminary investigation into differences in hops’ aroma attributes. Int. J. Food Sci. Technol. 53, 804–811 (2018).
De Keukeleire, J. et al. Relevance of organic farming and effect of climatological conditions on the formation of α-acids, β-acids, desmethylxanthohumol, and xanthohumol in hop (Humulus lupulus L.). J. Agric. Food Chem. 55, 61–66 (2007).
Matsui, H., Inui, T., Oka, K. & Fukui, N. The influence of pruning and harvest timing on hop aroma, cone appearance, and yield. Food Chem. 202, 15–22 (2016).
Pavlovic, V. et al. Environment and weather influence on quality and market value of hops. Plant Soil Environ. 58, 155–160 (2012).
Lafontaine, S. et al. Impact of harvest maturity on the aroma characteristics and chemistry of Cascade hops used for dry-hopping. Food Chem. 278, 228–239 (2019).
Van Simaeys, K. R. et al. Potential determinants of regional variation of three american aroma hops grown in the Willamette Valley, Oregon. J. Am. Soc. Brew. Chem. 80, 379–388 (2022).
Stark, C. & Gillespie, J. Suitability of New Zealand Cropping Regions to Support Hop Production. Lincoln University (2021). https://hapi.co.nz/wp-content/uploads/2021/08/Suitability-of-New-Zealand-cropping-regions-to-support-hop-production.pdf.
Pokorný, J., Pulkrábek, J., Štranc, P. & Bečka, D. Photosynthetic activity of selected genotypes of hops (Humulus lupulus L.) in critical periods for yield formation. Plant Soil Environ. 57, 264–270 (2011).
Potopová, V., Lhotka, O., Možný, M. & Musiolková, M. Vulnerability of hop‐yields due to compound drought and heat events over European key‐hop regions. Int. J. Climatol. 41, E2136–E2158 (2021).
Nesvadba, V., Hervert, J., Krofta, K. & Charvátová, J. Evaluation of Czech hop varieties in beer. Kvasny Prumysl 67, 529–536 (2021).
Hunter, J. J., Volschenk, C. G. & Zorer, R. Vineyard row orientation of Vitis vinifera L. cv. Shiraz/101-14 Mgt: climatic profiles and vine physiological status. Agric. Meteorol. 228–229, 104–119 (2016).
Neethling, E., Petitjean, T., Quénol, H. & Barbeau, G. Assessing local climate vulnerability and winegrowers’ adaptive processes in the context of climate change. Mitig. Adapt. Strateg. Glob. Change 22, 777–803 (2017).
Schindele, S. et al. Implementation of agrophotovoltaics: techno-economic analysis of the price-performance ratio and its policy implications. Appl. Energy 265, 114737 (2020).
Trommsdorff, M. et al. Combining food and energy production: design of an agrivoltaic system applied in arable and vegetable farming in Germany. Renew. Sustain. Energy Rev. 140, 110694 (2021).
Fandiño, M. et al. Assessing and modelling water use and the partition of evapotranspiration of irrigated hop (Humulus Lupulus), and relations of transpiration with hops yield and alpha-acids. Ind. Crops Prod. 77, 204–217 (2015).
Kolenc, Z. et al. Hop (Humulus lupulus L.) response mechanisms in drought stress: proteomic analysis with physiology. Plant Physiol. Biochem. 105, 67–78 (2016).
Nakawuka, P., Peters, T. R., Kenny, S. & Walsh, D. Effect of deficit irrigation on yield quantity and quality, water productivity and economic returns of four cultivars of hops in the Yakima Valley, Washington State. Ind. Crops Prod. 98, 82–92 (2017).
Potop, V., Možný, M. & Soukup, J. Drought evolution at various time scales in the lowland regions and their impact on vegetable crops in the Czech Republic. Agric. Meteorol. 156, 121–133 (2012).
Donner, P. et al. Influence of weather conditions, irrigation and plant age on yield and alpha-acids content of Czech hop (Humulus lupulus L.) cultivars. Plant Soil Environ. 66, 41–46 (2020).
Brant, V. et al. Distribution of root system of hop plants in hop gardens with regular rows cultivation. Plant Soil Environ. 66, 317–326 (2020).
Gent, D. H. et al. Delayed early season irrigation: impacts on hop yield and quality. J. Am. Soc. Brew. Chem. 80, 62–65 (2022).
FORHOPS. Asahi Europe & International. https://www.prochmel.cz/en/.
Bauerle, W. L. Intracanopy CO2 and light interactions on Humulus lupulus L. net canopy carbon gain under current and future atmospheric CO2 concentrations. Agric. Meteorol. 310, 108621 (2021).
Bauerle, W. L. Humulus lupulus L. Strobilus photosynthetic capacity and carbon assimilation. Plants 12, 1816 (2023).
Bauerle, W. L. Disentangling photoperiod from hop vernalization and dormancy for global production and speed breeding. Sci. Rep. 9, 16003 (2019).
Pörtner H.-O., et al. (eds.) IPCC, 2022: Climate change 2022: impacts, adaptation, and vulnerability. Contribution of Working Group II to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press. Cambridge University Press, Cambridge, UK and New York, NY, USA, 3056, https://doi.org/10.1017/9781009325844.
Lipper, L. et al. Climate-smart agriculture for food security. Nat. Clim. Change 4, 1068–1072 (2014).
Comi, M. Other agricultures of scale: social and environmental insights from Yakima Valley hop growers. J. Rural Stud. 80, 543–552 (2020).
BarthHaas. Hops are our passion. https://www.barthhaas.com/en/.
Pfeifroth, U., et al. Surface Radiation Data Set—Heliosat (SARAH)—Edition 2.1. Satell. Appl. Facil. Clim. Monit. https://doi.org/10.5676/EUM_SAF_CM/SARAH/V002_01 (2019).
Semenov, M. A., Donatelli, M., Stratonovitch, P., Chatzidaki, E. & Baruth, B. ELPIS: a dataset of local-scale daily climate scenarios for Europe. Clim. Res. 44, 3–15 (2010).
Mozny, M. et al. Climate-induced declines in the quality and quantity of European hops call for immediate adaptation measures (Data sources). Available at https://doi.org/10.6084/m9.figshare.23180342, Accesed 25 May 2023.
Acknowledgements
This work was supported by the project SustES—Adaptation strategies for sustainable ecosystem services and food security under adverse environmental conditions (CZ.02.1.01/0.0/0.0/16_019/0000797) to M.T., the Technology Agency of the Czech Republic (No. SS02030040, SS02030018) to M.M.
Author information
Authors and Affiliations
Contributions
The authors confirm contribution to the paper as follows: study conception and design: M.M., M.T., and U.B.; computation: M.M., V.V., and T.C.; data collection: V.V., M.S., L.H., and D.S. Analysis and interpretation of results: M.M., M.T., U.B., V.V., and Z.Z. All authors reviewed the results and approved the final version of the manuscript.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Communications thanks Hervé Quénol, and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. A peer review file is available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary information
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Mozny, M., Trnka, M., Vlach, V. et al. Climate-induced decline in the quality and quantity of European hops calls for immediate adaptation measures. Nat Commun 14, 6028 (2023). https://doi.org/10.1038/s41467-023-41474-5
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41467-023-41474-5