{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T16:52:18Z","timestamp":1785430338122,"version":"3.56.0"},"reference-count":186,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2026,6,12]],"date-time":"2026-06-12T00:00:00Z","timestamp":1781222400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Research Grants of the University of Thessaly"},{"id":[{"id":"https:\/\/ror.org\/04v4g9h31","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Heat stress (HS) is at the top of the challenges facing modern dairy production, with annual losses according to global projections, under high-emission scenarios, reaching US$14.7\u201340.0 billion by the end of the century. This review emphasizes three interconnected topics that account for most of the proportion of the productive and reproductive losses during HS. First, the physiological consequences of HS are reviewed, with emphasis on the pair-fed thermal neutral (PFTN) paradigm, which established that reduced dry matter intake (DMI) accounts for only 35\u201350% of the observed milk yield decline, with the remainder arising from tissue-level effects of hyperthermia on mammary function, metabolism, and reproductive performance. Second, HS-induced microbiome disruption is examined as an active pathophysiological amplifier, whereby rumen dysbiosis compromises intestinal barrier integrity and drives systemic endotoxaemia, chronically amplifying the immune suppression already imposed by the thermal insult. Third, we focus on the integration of multi-omics platforms as a management approach, since single-omics analyses capture only a fraction of the biological complexity underlying the HS response. As the available datasets expand in coverage and scale, their integration through AI-driven analytical frameworks has the potential to substantially advance beyond the current fragmented picture, progressively building toward a systems-level model of thermal stress. Evidence-based mitigation strategies spanning environmental cooling, targeted nutritional supplementation, and genomic selection are critically evaluated within this framework, with emphasis on equity of access to evidence-based solutions across global dairy production systems.<\/jats:p>","DOI":"10.3390\/biology15120918","type":"journal-article","created":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T00:49:42Z","timestamp":1781743782000},"page":"918","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Consequences of Heat Stress on Physiology, Microbiome Dynamics, and Multi-Omics in Dairy Cows: More than Meets the Eye"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1842-5432","authenticated-orcid":false,"given":"Themistoklis","family":"Giannoulis","sequence":"first","affiliation":[{"name":"Department of Animal Science, University of Thessaly, 41223 Larissa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4027-2660","authenticated-orcid":false,"given":"Eleni","family":"Dovolou","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of Thessaly, 41223 Larissa, Greece"},{"name":"Department of Obstetrics & Reproduction, Faculty of Veterinary Science, University of Thessaly, 43100 Karditsa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6868-8927","authenticated-orcid":false,"given":"Zissis","family":"Mamuris","sequence":"additional","affiliation":[{"name":"Laboratory of Genetics, Comparative and Evolutionary Biology, Department of Biochemistry and Biotechnology, University of Thessaly, 41500 Larissa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Georgios S.","family":"Amiridis","sequence":"additional","affiliation":[{"name":"Department of Obstetrics & Reproduction, Faculty of Veterinary Science, University of Thessaly, 43100 Karditsa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,6,12]]},"reference":[{"key":"ref_1","unstructured":"IPCC (2023). Climate Change 2023: Synthesis Report, IPCC. Contribution of Working Groups I, II and III to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"E52","DOI":"10.3168\/jds.S0022-0302(03)74040-5","article-title":"Economic losses from heat stress by US livestock industries","volume":"86","author":"Cobanov","year":"2003","journal-title":"J. Dairy Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"e192","DOI":"10.1016\/S2542-5196(22)00002-X","article-title":"Impacts of heat stress on global cattle production during the 21st century: A modelling study","volume":"6","author":"Thornton","year":"2022","journal-title":"Lancet Planet. Health"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"859","DOI":"10.5194\/esd-10-859-2019","article-title":"Heat stress risk in European dairy cattle husbandry under different climate change scenarios","volume":"10","author":"Hempel","year":"2019","journal-title":"Earth Syst. Dynam."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/S0301-6226(01)00330-X","article-title":"Heat stress in lactating dairy cows: A review","volume":"77","author":"Kadzere","year":"2002","journal-title":"Livest. Prod. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"10367","DOI":"10.3168\/jds.2017-13676","article-title":"A 100-year review: Stress physiology including heat stress","volume":"100","author":"Collier","year":"2017","journal-title":"J. Dairy Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1093\/af\/vfy039","article-title":"Climate change: Impact on livestock and how can we adapt","volume":"9","author":"Bernabucci","year":"2019","journal-title":"Anim. Front."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.smallrumres.2014.11.005","article-title":"Impact of climate change on the dairy industry in temperate zones: Predictions on the overall negative impact and on the positive role of dairy goats in adapting the industry to temperature and dryness stresses","volume":"123","author":"Silanikove","year":"2015","journal-title":"Small Rumin. Res."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"s431","DOI":"10.1017\/S1751731118001945","article-title":"Review: Adaptation of animals to heat stress","volume":"12","author":"Sejian","year":"2018","journal-title":"Animal"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Cheruiyot, E.K., Nguyen, T.T.T., Haile-Mariam, M., Cocks, B.G., Abdelsayed, M., and Pryce, J.E. (2022). Improving genomic selection for heat tolerance in dairy cattle: Current opportunities and future directions. Front. Genet., 13.","DOI":"10.3389\/fgene.2022.894067"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Cartwright, S.L., Verschoor, C.P., and Karrow, N.A. (2023). Invited Review: Impact of heat stress on dairy cattle and selection strategies for thermotolerance. Front. Vet. Sci., 10.","DOI":"10.3389\/fvets.2023.1198697"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"3207","DOI":"10.3168\/jds.2023-24059","article-title":"Effects of heat stress on feed intake, milk yield, milk composition, and feed efficiency in dairy cows: A meta-analysis","volume":"107","author":"Chen","year":"2024","journal-title":"J. Dairy Sci."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1016\/j.theriogenology.2020.02.048","article-title":"Impact of heat stress on lactational performance of dairy cows","volume":"150","author":"Tao","year":"2020","journal-title":"Theriogenology"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1017\/S0021859600072117","article-title":"Diurnal patterns of heat production and heart rate under thermoneutral conditions in Holstein Friesian cows differing in milk production","volume":"114","author":"Purwanto","year":"1990","journal-title":"J. Agric. Sci."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"3798","DOI":"10.3168\/jds.2015-10212","article-title":"Modeling heat stress under different environmental conditions","volume":"99","author":"Carabano","year":"2016","journal-title":"J. Dairy Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1986","DOI":"10.3168\/jds.2008-1641","article-title":"Effects of heat stress and plane of nutrition on lactating Holstein cows: I. Production, metabolism, and aspects of circulating somatotropin","volume":"92","author":"Rhoads","year":"2009","journal-title":"J. Dairy Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"644","DOI":"10.3168\/jds.2009-2295","article-title":"Effects of heat stress on energetic metabolism in lactating Holstein cows","volume":"93","author":"Wheelock","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"5040","DOI":"10.3168\/jds.2016-11913","article-title":"The effects of heat stress on protein metabolism in lactating Holstein cows","volume":"100","author":"Gao","year":"2017","journal-title":"J. Dairy Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"257","DOI":"10.2527\/jas.52339","article-title":"Heat stress reduces barrier function and alters intestinal metabolism in growing pigs","volume":"90","author":"Pearce","year":"2012","journal-title":"J. Anim. Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"7842","DOI":"10.3168\/jds.2022-21820","article-title":"Heat stress develops with increased total-tract gut permeability, and dietary organic acid and pure botanical supplementation partly restores lactation performance in Holstein dairy cows","volume":"105","author":"Fontoura","year":"2022","journal-title":"J. Dairy Sci."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"10333","DOI":"10.1073\/pnas.1820130116","article-title":"Heat stress directly impairs gut integrity and recruits distinct immune cell populations into the bovine intestine","volume":"116","author":"Koch","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Oliveira, C.P., Sousa, F.C.d., da Silva, A.L., Schultz, \u00c9.B., Valderrama Londo\u00f1o, R.I., and de Souza, P.A.R. (2025). Heat stress in dairy cows: Impacts, identification, and mitigation strategies\u2014A review. Animals, 15.","DOI":"10.3390\/ani15020249"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2131","DOI":"10.3168\/jds.S0022-0302(03)73803-X","article-title":"Effects of heat-stress on production in dairy cattle","volume":"86","author":"West","year":"2003","journal-title":"J. Dairy Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1093\/af\/vfy031","article-title":"Heat stress: Physiology of acclimation and adaptation","volume":"9","author":"Collier","year":"2019","journal-title":"Anim. Front."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1017\/S1751731111002448","article-title":"Adaptation to hot climate and strategies to alleviate heat stress in livestock production","volume":"6","author":"Renaudeau","year":"2012","journal-title":"Animal"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0301-6226(00)00162-7","article-title":"Effects of heat stress on the welfare of extensively managed domestic ruminants","volume":"67","author":"Silanikove","year":"2000","journal-title":"Livest. Prod. Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"711","DOI":"10.13031\/2013.34325","article-title":"Black globe-humidity index (BGHI) as comfort equation for dairy cows","volume":"24","author":"Buffington","year":"1981","journal-title":"Trans. ASAE"},{"key":"ref_28","unstructured":"Zimbelman, R.B., Rhoads, R.P., Rhoads, M.L., and Baumgard, L.H. (2011, January 9\u201311). A re-evaluation of the impact of temperature humidity index (THI) and black globe humidity index (BGHI) on milk production in high producing dairy cows. Proceedings of the Western Dairy Management Conference, Reno, NV, USA."},{"key":"ref_29","first-page":"4837","article-title":"The effects of heat stress in Italian Holstein dairy cattle","volume":"93","author":"Bernabucci","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1377","DOI":"10.2527\/2005.8361377x","article-title":"Estimates of heat stress relief needs for Holstein dairy cows","volume":"83","author":"Berman","year":"2005","journal-title":"J. Anim. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1811","DOI":"10.1007\/s00484-022-02321-2","article-title":"Thermodynamic assessment of heat stress in dairy cattle: Lessons from human biometeorology","volume":"66","author":"Foroushani","year":"2022","journal-title":"Int. J. Biometeorol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Gunn, K., Holly, M., Veith, T., Buda, A., Prasad, R., Rotz, C., Soder, K., and Stoner, A. (2019). Projected heat stress challenges and abatement opportunities for U.S. milk production. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0214665"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Yan, G., Li, H., and Shi, Z. (2021). Evaluation of Thermal Indices as the Indicators of Heat Stress in Dairy Cows in a Temperate Climate. Animals, 11.","DOI":"10.3390\/ani11082459"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Xiong, Z., Li, L., Ouyang, K., Qu, M., and Qiu, Q. (2025). Deciphering Heat Stress Mechanisms and Developing Mitigation Strategies in Dairy Cattle: A Multi-Omics Perspective. Agriculture, 15.","DOI":"10.3390\/agriculture15141477"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Lezama-Garc\u00eda, K., Mota-Rojas, D., Pereira, A.M.F., Mart\u00ednez-Burnes, J., Ghezzi, M., Dom\u00ednguez, A., G\u00f3mez, J., de Mira Geraldo, A., Lendez, P., and Hern\u00e1ndez-\u00c1valos, I. (2022). Transient Receptor Potential (TRP) and thermoregulation in animals: Structural biology and neurophysiological aspects. Animals, 12.","DOI":"10.3390\/ani12010106"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1146\/annurev-animal-031412-103644","article-title":"Effects of heat stress on postabsorptive metabolism and energetics","volume":"1","author":"Baumgard","year":"2013","journal-title":"Annu. Rev. Anim. Biosci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2035","DOI":"10.3168\/jds.2022-22489","article-title":"Evaporative Water Loss from Dairy Cows in Climate-Controlled Respiration Chambers","volume":"106","author":"Zhou","year":"2023","journal-title":"J. Dairy Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"5642","DOI":"10.3168\/jds.2017-13727","article-title":"Symposium review: The influences of heat stress on bovine mammary gland function","volume":"101","author":"Tao","year":"2018","journal-title":"J. Dairy Sci."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Fu, L., Zhang, L., Liu, L., Yang, H., Zhou, P., Song, F., Dong, G., Chen, J., Wang, G., and Dong, X. (2021). Effect of Heat Stress on Bovine Mammary Cellular Metabolites and Gene Transcription Related to Amino Acid Metabolism, Amino Acid Transportation and Mammalian Target of Rapamycin (mTOR) Signaling. Animals, 11.","DOI":"10.3390\/ani11113153"},{"key":"ref_40","first-page":"342","article-title":"Early-life heat stress exposure impacts dairy calf feeding and thermoregulatory behavior","volume":"2","author":"Gingerich","year":"2021","journal-title":"JDS Commun."},{"key":"ref_41","first-page":"119","article-title":"Nutrient supply to high-yielding cows","volume":"26","author":"Beever","year":"2001","journal-title":"Br. Soc. Anim. Sci."},{"key":"ref_42","first-page":"57","article-title":"Physiological constraints to milk production: Factors which determine nutrient partitioning, lactation persistency, and mobilization of body reserves","volume":"34","author":"Chilliard","year":"1992","journal-title":"World Rev. Anim. Prod."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.livsci.2010.02.011","article-title":"Effects of climate changes on animal production and sustainability of livestock systems","volume":"130","author":"Nardone","year":"2010","journal-title":"Livest. Sci."},{"key":"ref_44","unstructured":"Collier, R.J., Baumgard, L.H., Lock, A.L., and Bauman, D.E. (2005). Physiological limitations, nutrient partitioning. Yields of Farmed Species: Constraints and Opportunities in the 21st Century, Nottingham University Press."},{"key":"ref_45","first-page":"53","article-title":"Body condition score, metabolic status and milk production of early lactating dairy cows exposed to warm or cold environments","volume":"22","author":"Lacetera","year":"1996","journal-title":"Zootec. Nutr. Anim."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.3168\/jds.S0022-0302(05)72878-2","article-title":"Influence of body condition score on relationships between metabolic status and oxidative stress in periparturient dairy cows","volume":"88","author":"Bernabucci","year":"2005","journal-title":"J. Dairy Sci."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"267","DOI":"10.3945\/an.112.003376","article-title":"Nutritional interventions to alleviate the negative consequences of heat stress","volume":"4","author":"Rhoads","year":"2013","journal-title":"Adv. Nutr."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Sammad, A., Wang, Y.J., Umer, S., Lirong, H., Khan, I., Khan, A., Ahmad, B., and Wang, Y. (2020). Nutritional Physiology and Biochemistry of Dairy Cattle under the Influence of Heat Stress: Consequences and Opportunities. Animals, 10.","DOI":"10.3390\/ani10050793"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"170","DOI":"10.3168\/jds.2009-2469","article-title":"Effects of heat stress and nutrition on lactating Holstein cows: II. Aspects of hepatic growth hormone responsiveness","volume":"93","author":"Rhoads","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.jprot.2017.02.011","article-title":"Seasonal heat stress affects adipose tissue proteome toward enrichment of the Nrf2-mediated oxidative stress response in late-pregnant dairy cows","volume":"158","author":"Zachut","year":"2017","journal-title":"J. Proteom."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Gupta, S., Sharma, A., Joy, A., Dunshea, F.R., and Chauhan, S.S. (2023). The Impact of Heat Stress on Immune Status of Dairy Cattle and Strategies to Ameliorate the Negative Effects. Animals, 13.","DOI":"10.3390\/ani13010107"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1016\/j.rvsc.2019.08.011","article-title":"The impact of heat stress on the immune system in dairy cattle: A review","volume":"126","author":"Bagath","year":"2019","journal-title":"Res. Vet. Sci."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Dahl, G.E., Tao, S., and Laporta, J. (2020). Heat stress impacts immune status in cows across the life cycle. Front. Vet. Sci., 7.","DOI":"10.3389\/fvets.2020.00116"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2173","DOI":"10.3168\/jds.S0022-0302(02)74296-3","article-title":"Markers of oxidative status in plasma and erythrocytes of transition dairy cows during hot season","volume":"85","author":"Bernabucci","year":"2002","journal-title":"J. Dairy Sci."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Kim, H., Jo, J.-H., Lee, H.-G., Park, W., Lee, H.-K., Park, J.-E., and Shin, D. (2024). Inflammatory Response in Dairy Cows Caused by Heat Stress and Biological Mechanisms for Maintaining Homeostasis. PLoS ONE, 19.","DOI":"10.1371\/journal.pone.0300719"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"8645","DOI":"10.3168\/jds.2017-12651","article-title":"Invited review: Effects of heat stress on dairy cattle welfare","volume":"100","author":"Polsky","year":"2017","journal-title":"J. Dairy Sci."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"3008","DOI":"10.3168\/jds.2022-22520","article-title":"Mild heat stress-induced adaptive immune response in blood mononuclear cells and leukocytes from mesenteric lymph nodes of primiparous lactating Holstein cows","volume":"106","author":"Koch","year":"2023","journal-title":"J. Dairy Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"118","DOI":"10.3168\/jds.2013-7704","article-title":"Effect of core body temperature, time of day, and climate conditions on behavioral patterns of lactating dairy cows experiencing mild to moderate heat stress","volume":"98","author":"Allen","year":"2015","journal-title":"J. Dairy Sci."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"4765","DOI":"10.2527\/jas.2011-5064","article-title":"Relationships between rumination time, metabolic conditions, and health status in dairy cows during the first weeks of lactation","volume":"90","author":"Soriani","year":"2012","journal-title":"J. Anim. Sci."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"9057","DOI":"10.3168\/jds.2016-11203","article-title":"Peripartal Rumination Dynamics and Health Status in Cows Calving in Hot and Cool Seasons","volume":"99","author":"Paudyal","year":"2016","journal-title":"J. Dairy Sci."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/j.theriogenology.2022.08.001","article-title":"Effect of calving season on metritis incidence and bacterial content of the vagina in dairy cows","volume":"191","author":"Molinari","year":"2022","journal-title":"Theriogenology"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1464","DOI":"10.3168\/jds.2022-22405","article-title":"Prepartum heat stress increases postpartum inflammatory responses in blood of lactating dairy cows","volume":"106","author":"Molinari","year":"2023","journal-title":"J. Dairy Sci."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"6751","DOI":"10.3168\/jds.2019-17929","article-title":"Invited Review: Physiological and Behavioral Effects of Heat Stress in Dairy Cows","volume":"103","author":"Becker","year":"2020","journal-title":"J. Dairy Sci."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1674","DOI":"10.3168\/jds.2006-634","article-title":"The effect of heat stress and lameness on time budgets of lactating dairy cows","volume":"90","author":"Cook","year":"2007","journal-title":"J. Dairy Sci."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Gal\u00e1n, E., Llonch, P., Villagr\u00e1, A., Levit, H., Pinto, S., and Del Prado, A. (2018). A systematic review of non-productivity-related animal-based indicators of heat stress resilience in dairy cattle. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0206520"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"3716","DOI":"10.3168\/jds.2021-21080","article-title":"Invited review: Lethal heat stress\u2014The putative pathophysiology of a deadly disorder in dairy cattle","volume":"105","author":"Burhans","year":"2022","journal-title":"J. Dairy Sci."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"3781","DOI":"10.3168\/jds.2009-2127","article-title":"Seasonal pattern of mortality and relationships between mortality and temperature-humidity index in dairy cows","volume":"92","author":"Vitali","year":"2009","journal-title":"J. Dairy Sci."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Joo, S.S., Lee, S.J., Park, D.S., Kim, D.H., Gu, B.-H., Park, Y.J., Rim, C.Y., Kim, M., and Kim, E.T. (2021). Changes in Blood Metabolites and Immune Cells in Holstein and Jersey Dairy Cows by Heat Stress. Animals, 11.","DOI":"10.3390\/ani11040974"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Velayudhan, S.M., Br\u00fcgemann, K., Alam, S., Yin, T., Devaraj, C., Sejian, V., Schlecht, E., and K\u00f6nig, S. (2023). Molecular, Physiological and Hematological Responses of Crossbred Dairy Cattle in a Tropical Savanna Climate. Biology, 12.","DOI":"10.3390\/biology12010026"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1692","DOI":"10.2460\/ajvr.1991.52.10.1692","article-title":"Modulation of function of bovine polymorphonuclear leukocytes and lymphocytes by high temperature in vitro and in vivo","volume":"52","author":"Elvinger","year":"1991","journal-title":"Am. J. Vet. Res."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"4606","DOI":"10.3168\/jds.S0022-0302(06)72510-3","article-title":"Heat stress elicits different responses in peripheral blood mononuclear cells from brown Swiss and Holstein cows","volume":"89","author":"Lacetera","year":"2006","journal-title":"J. Dairy Sci."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"445","DOI":"10.3168\/jds.2007-0540","article-title":"Invited review: Genes involved in the bovine heat stress response","volume":"91","author":"Collier","year":"2008","journal-title":"J. Dairy Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1093\/af\/vfy027","article-title":"Impact of heat stress on cow reproduction and fertility","volume":"9","author":"Wolfenson","year":"2019","journal-title":"Anim. Front."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1093\/jas\/skaa139","article-title":"Influence of heat stress on reproduction in dairy cows\u2014Physiological and practical aspects","volume":"98","author":"Roth","year":"2020","journal-title":"J. Anim. Sci."},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Dovolou, E., Giannoulis, T., Nanas, I., and Amiridis, G.S. (2023). Heat stress: A serious disruptor of the reproductive physiology of dairy cows. Animals, 13.","DOI":"10.3390\/ani13111846"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"978","DOI":"10.1111\/jpn.13257","article-title":"Dairy cow reproduction under the influence of heat stress","volume":"104","author":"Sammad","year":"2020","journal-title":"J. Anim. Physiol. Anim. Nutr."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1139","DOI":"10.1016\/S0093-691X(03)00126-2","article-title":"Heat stress and seasonal effects on reproduction in the dairy cow\u2014A review","volume":"60","author":"Scaramuzzi","year":"2003","journal-title":"Theriogenology"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1774","DOI":"10.1111\/rda.13840","article-title":"A study on stress response and fertility parameters in phenotypically thermotolerant and thermosensitive dairy cows during summer heat stress","volume":"55","author":"Nanas","year":"2020","journal-title":"Reprod. Domest. Anim."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"103564","DOI":"10.1016\/j.jtherbio.2023.103564","article-title":"Inflammatory responses of bovine endometrial epithelial cells are increased under in vitro heat stress conditions","volume":"114","author":"Molinari","year":"2023","journal-title":"J. Therm. Biol."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"G\u00f3mez-Guzm\u00e1n, J.A., Parra-Bracamonte, G.M., and Velazquez, M.A. (2024). Impact of heat stress on oocyte developmental competence and pre-implantation embryo viability in cattle. Animals, 14.","DOI":"10.3390\/ani14152280"},{"key":"ref_81","unstructured":"Carreira, R.P. (2017). Mitigation of the Heat Stress Impact in Livestock Reproduction, IntechOpen."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1379","DOI":"10.1016\/j.theriogenology.2007.02.009","article-title":"Climate factors affecting conception rate of high producing dairy cows in northeastern Spain","volume":"67","author":"Santolaria","year":"2007","journal-title":"Theriogenology"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1146\/annurev-animal-022516-022849","article-title":"Effect of heat stress on reproduction in dairy cows: Insights into the cellular and molecular responses of the oocyte","volume":"5","author":"Roth","year":"2017","journal-title":"Annu. Rev. Anim. Biosci."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"3642","DOI":"10.3168\/jds.2017-13389","article-title":"Reduction in oocyte developmental competence by stress is associated with alterations in mitochondrial function","volume":"101","author":"Roth","year":"2018","journal-title":"J. Dairy Sci."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1016\/j.theriogenology.2020.06.039","article-title":"Short term temperature elevation during IVM affects embryo yield and alters gene expression pattern in oocytes, cumulus cells and blastocysts in cattle","volume":"156","author":"Stamperna","year":"2020","journal-title":"Theriogenology"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1302","DOI":"10.1111\/rda.13993","article-title":"Developmental competence of heat stressed oocytes from Holstein and Limousine cows matured in vitro","volume":"56","author":"Stamperna","year":"2021","journal-title":"Reprod. Domest. Anim."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1002\/(SICI)1098-2795(199702)46:2<138::AID-MRD4>3.0.CO;2-R","article-title":"Differential responses of bovine oocytes and pre-implantation embryos to heat shock","volume":"46","author":"Edwards","year":"1997","journal-title":"Mol. Reprod. Dev."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"S242","DOI":"10.1016\/j.theriogenology.2007.04.008","article-title":"Exploitation of genetic and physiological determinants of embryonic resistance to elevated temperature to improve embryonic survival in dairy cattle under heat stress","volume":"68","author":"Hansen","year":"2007","journal-title":"Theriogenology"},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Mi\u0119tkiewska, K., Kordowitzki, P., and Pareek, C.S. (2022). Effects of Heat Stress on Bovine Oocytes and Early Embryonic Development\u2014An Update. Cells, 11.","DOI":"10.3390\/cells11244073"},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Feng, X., Li, C., Zhang, H., Zhang, P., Shahzad, M., Du, W., and Zhao, X. (2024). Heat-Stress Impacts on Developing Bovine Oocytes: Unraveling Epigenetic Changes, Oxidative Stress, and Developmental Resilience. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25094808"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1016\/S0378-4320(00)00102-0","article-title":"Impaired reproduction in heat-stressed cattle: Basic and applied aspects","volume":"60\u201361","author":"Wolfenson","year":"2000","journal-title":"Anim. Reprod. Sci."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.anireprosci.2009.04.013","article-title":"Biochemical changes in the follicular fluid of the dominant follicle of high producing dairy cows exposed to heat stress early post-partum","volume":"117","author":"Leroy","year":"2010","journal-title":"Anim. Reprod. Sci."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"102951","DOI":"10.1016\/j.jtherbio.2021.102951","article-title":"Early embryo losses, progesterone and pregnancy associated glycoproteins levels during summer heat stress in dairy cows","volume":"98","author":"Nanas","year":"2021","journal-title":"J. Therm. Biol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.theriogenology.2022.04.011","article-title":"Oviductal epithelial cells transcriptome and extracellular vesicles characterization during thermoneutral and heat stress conditions in dairy cows","volume":"187","author":"Stamperna","year":"2022","journal-title":"Theriogenology"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"6308","DOI":"10.3168\/jds.2023-24606","article-title":"Programming effects of intrauterine hyperthermia on adrenal gland development","volume":"107","author":"Guadagnin","year":"2024","journal-title":"J. Dairy Sci."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"2976","DOI":"10.3168\/jds.2016-11993","article-title":"In utero exposure to heat stress during late gestation has prolonged effects on the activity patterns and growth of dairy calves","volume":"100","author":"Laporta","year":"2017","journal-title":"J. Dairy Sci."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"165","DOI":"10.3168\/jdsc.2020-0069","article-title":"Effect of late-gestation heat stress in nulliparous heifers on postnatal growth, passive transfer of immunoglobulin G, and thermoregulation of their calves","volume":"2","author":"Davidson","year":"2021","journal-title":"JDS Commun."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Dado-Senn, B., Skibiel, A.L., Fabris, T.F., Dahl, G.E., and Laporta, J. (2019). Dry period heat stress induces microstructural changes in the lactating mammary gland and alters the DNA methylome of the mammary epithelium. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0222120"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"7555","DOI":"10.3168\/jds.2020-18154","article-title":"Late-Gestation Heat Stress Impairs Daughter and Granddaughter Lifetime Performance","volume":"103","author":"Laporta","year":"2020","journal-title":"J. Dairy Sci."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1016\/j.theriogenology.2020.03.011","article-title":"Late gestation heat stress in dairy cows: Effects on dam and daughter","volume":"150","author":"Ouellet","year":"2020","journal-title":"Theriogenology"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"6426","DOI":"10.3168\/jds.2013-7891","article-title":"Effect of heat stress during late gestation on immune function and growth performance of calves: Isolation of altered colostral and calf factors","volume":"97","author":"Monteiro","year":"2014","journal-title":"J. Dairy Sci."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"3193","DOI":"10.3168\/jds.2015-9990","article-title":"Effects of late-gestation heat stress on immunity and performance of calves","volume":"99","author":"Dahl","year":"2016","journal-title":"J. Dairy Sci."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"4079","DOI":"10.3168\/jds.2012-6278","article-title":"Invited review: Heat stress effects during late gestation on dry cows and their calves","volume":"96","author":"Tao","year":"2013","journal-title":"J. Dairy Sci."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"628","DOI":"10.15232\/S1080-7446(15)30421-6","article-title":"Dry-period seasonal effects on the subsequent lactation","volume":"28","author":"Thompson","year":"2012","journal-title":"Prof. Anim. Sci."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"4390","DOI":"10.3168\/jds.2024-25975","article-title":"Preweaning heat stress alters liver transcriptome and DNA methylation in dairy calves","volume":"108","author":"Laporta","year":"2025","journal-title":"J. Dairy Sci."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"17021","DOI":"10.1038\/s41598-025-01082-3","article-title":"Heat stress induces specific methylation, transcriptomic and metabolic patterns in dairy cows and their female progeny","volume":"15","author":"Halli","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"5976","DOI":"10.3168\/jds.2011-4329","article-title":"Effect of Heat Stress during the Dry Period on Mammary Gland Development","volume":"94","author":"Tao","year":"2011","journal-title":"J. Dairy Sci."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"4822","DOI":"10.3168\/jds.2019-17926","article-title":"Pre- and postnatal heat stress abatement affects dairy calf thermoregulation and performance","volume":"103","author":"Field","year":"2020","journal-title":"J. Dairy Sci."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"10695","DOI":"10.3168\/jds.2019-16718","article-title":"Invited review: Management strategies capable of improving the reproductive performance of heat-stressed dairy cattle","volume":"102","author":"Fausnacht","year":"2019","journal-title":"J. Dairy Sci."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1016\/j.theriogenology.2015.04.021","article-title":"Seasonal heat stress: Clinical implications and hormone treatments for the fertility of dairy cows","volume":"84","year":"2015","journal-title":"Theriogenology"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"S36","DOI":"10.1262\/jrd.1056S36","article-title":"Management of heat stress to improve fertility in dairy cows in Israel","volume":"56","author":"Flamenbaum","year":"2010","journal-title":"J. Reprod. Dev."},{"key":"ref_112","doi-asserted-by":"crossref","unstructured":"He, Y., Maltecca, C., and Tiezzi, F. (2021). Potential use of gut microbiota composition as a biomarker of heat stress in monogastric species: A review. Animals, 11.","DOI":"10.3390\/ani11061833"},{"key":"ref_113","doi-asserted-by":"crossref","unstructured":"Kim, S.C., Ramos, S.C., Valencia, R.A., Cho, Y., and Lee, S.S. (2022). Heat stress: Effects on rumen microbes and host physiology, and strategies to alleviate the negative impacts on lactating dairy cows. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.804562"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"12824","DOI":"10.3168\/jds.2025-26997","article-title":"Integrative Blood Transcriptomic and Metabolomic Profiling Reveals Biomarkers of Natural Heat Tolerance in Holstein Cows","volume":"108","author":"Li","year":"2025","journal-title":"J. Dairy Sci."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"103562","DOI":"10.1016\/j.jtherbio.2023.103562","article-title":"Heat stress promotes adaptive physiological responses and alters mRNA expression of ruminal epithelium markers in Bos taurus indicus cattle fed low- or high-energy diets","volume":"114","author":"Meneses","year":"2023","journal-title":"J. Therm. Biol."},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Zhao, S., Min, L., Zheng, N., and Wang, J. (2019). Effect of Heat Stress on Bacterial Composition and Metabolism in the Rumen of Lactating Dairy Cows. Animals, 9.","DOI":"10.3390\/ani9110925"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"279","DOI":"10.3168\/jds.2009-2207","article-title":"pH dynamics and bacterial community composition in the rumen of lactating dairy cows","volume":"93","author":"Palmonari","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_118","doi-asserted-by":"crossref","unstructured":"Guo, Z., Gao, S., Ding, J., He, J., Ma, L., and Bu, D. (2021). Effects of Heat Stress on the Ruminal Epithelial Barrier of Dairy Cows Revealed by Micromorphological Observation and Transcriptomic Analysis. Front. Genet., 12.","DOI":"10.3389\/fgene.2021.768209"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"644","DOI":"10.1038\/s41597-022-01777-6","article-title":"Effects of heat stress on 16S rDNA, metagenome and metabolome in Holstein cows at different growth stages","volume":"9","author":"Feng","year":"2022","journal-title":"Sci. Data"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"e0331223","DOI":"10.1128\/spectrum.03312-23","article-title":"Altered rumen microbiome and correlations of the metabolome in heat-stressed dairy cows at different growth stages","volume":"11","author":"Feng","year":"2023","journal-title":"Microbiol. Spectr."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"5162","DOI":"10.3168\/jds.2023-24242","article-title":"Heat stress and feeding effects on the mucosa-associated and digesta microbiome and their relationship to plasma and digesta fluid metabolites in the jejunum of dairy cows","volume":"107","author":"Koch","year":"2024","journal-title":"J. Dairy Sci."},{"key":"ref_122","doi-asserted-by":"crossref","unstructured":"Wang, Z., Guo, M., Liang, Y., Zhou, F., Zhang, H., Li, M., Yang, Z., Karrow, N., and Mao, Y. (2025). Breed-specific responses and ruminal microbiome shifts in dairy cows under heat stress. Animals, 15.","DOI":"10.3390\/ani15060817"},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"Oikonomou, G., Machado, V.S., Santisteban, C., Schukken, Y.H., and Bicalho, R.C. (2012). Microbial diversity of bovine mastitic milk as described by pyrosequencing of metagenomic 16s rDNA. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0047671"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"664","DOI":"10.1111\/1574-6976.12030","article-title":"The Complex Microbiota of Raw Milk","volume":"37","author":"Quigley","year":"2013","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"11639","DOI":"10.3168\/jds.2024-24976","article-title":"Heat stress differentially alters the milk microbiota of Holstein Friesian and Brown Swiss dairy cows","volume":"107","author":"Ceciliani","year":"2024","journal-title":"J. Dairy Sci."},{"key":"ref_126","doi-asserted-by":"crossref","unstructured":"Kotsampasi, B., Karatzia, M.A., Tsiokos, D., and Chadio, S. (2024). Nutritional strategies to alleviate stress and improve welfare in dairy ruminants. Animals, 14.","DOI":"10.3390\/ani14172573"},{"key":"ref_127","first-page":"G1195","article-title":"Splanchnic tissues undergo hypoxic stress during whole body hyperthermia","volume":"276","author":"Hall","year":"1999","journal-title":"Am. J. Physiol."},{"key":"ref_128","first-page":"G204","article-title":"Physiologically relevant increase in temperature causes an increase in intestinal epithelial tight junction permeability","volume":"290","author":"Dokladny","year":"2006","journal-title":"Am. J. Physiol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"1452","DOI":"10.2527\/jas.2011-4627","article-title":"Endotoxin, inflammation, and intestinal function in livestock","volume":"90","author":"Mani","year":"2012","journal-title":"J. Anim. Sci."},{"key":"ref_130","doi-asserted-by":"crossref","unstructured":"Scerri, T.M., Lomax, S., and Clark, C.E.F. (2023). Bovine heat stress management: Current amelioration approaches and the case for a novel mitogenomic strategy. Front. Anim. Sci., 4.","DOI":"10.3389\/fanim.2023.1169743"},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"1149","DOI":"10.1007\/s00484-016-1283-z","article-title":"Metabolic responses and \u201comics\u201d technologies for elucidating the effects of heat stress in dairy cows","volume":"61","author":"Min","year":"2017","journal-title":"Int. J. Biometeorol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1093\/af\/vfy033","article-title":"Selecting for heat tolerance","volume":"9","author":"Carabano","year":"2019","journal-title":"Anim. Front."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"13512","DOI":"10.3168\/jds.2025-26818","article-title":"Development of Genomic Predictions for Heat Tolerance in US Holstein Cattle","volume":"108","author":"Vukasinovic","year":"2025","journal-title":"J. Dairy Sci."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"16619","DOI":"10.1038\/s41598-021-95816-8","article-title":"New loci and neuronal pathways for resilience to heat stress in cattle","volume":"11","author":"Cheruiyot","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"2849","DOI":"10.3168\/jds.2015-9685","article-title":"Genomic selection for tolerance to heat stress in Australian dairy cattle","volume":"99","author":"Nguyen","year":"2016","journal-title":"J. Dairy Sci."},{"key":"ref_136","doi-asserted-by":"crossref","unstructured":"Worku, D., Hussen, J., De Matteis, G., Schusser, B., and Alhussien, M.N. (2023). Candidate genes associated with heat stress and breeding strategies to relieve its effects in dairy cattle: A deeper insight into the genetic architecture and immune response to heat stress. Front. Vet. Sci., 10.","DOI":"10.3389\/fvets.2023.1151241"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1007\/s12272-021-01336-4","article-title":"Recent advances in CRISPR technologies for genome editing","volume":"44","author":"Song","year":"2021","journal-title":"Arch. Pharm. Res."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"5508","DOI":"10.3168\/jds.2014-8087","article-title":"The SLICK hair locus derived from Senepol cattle confers thermotolerance to intensively managed lactating Holstein cows","volume":"97","author":"Dikmen","year":"2014","journal-title":"J. Dairy Sci."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1096\/fba.2024-00029","article-title":"Consequences of gene editing of PRLR on thermotolerance, growth, and male reproduction in cattle","volume":"6","author":"Cuellar","year":"2024","journal-title":"FASEB Bioadv."},{"key":"ref_140","unstructured":"US Food and Drug Administration (2022). FDA Issues Its First Low Risk Determination for a Beef Cattle Breed Developed Using Gene Editing."},{"key":"ref_141","unstructured":"Hansen, P.J. (2002, January 19\u201323). Countering heat stress in cattle: Strategies based on temperature regulation, the biology of heat tolerance, and molecular biology. Proceedings of the 7th World Congress on Genetics Applied to Livestock Production, Montpellier, France."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1111\/rda.14274","article-title":"Genetic diversity and thermotolerance in Holstein cows: Pathway analysis and marker development using whole-genome sequencing","volume":"58","author":"Kalemkeridou","year":"2022","journal-title":"Reprod. Domest. Anim."},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Li, G., Yu, X., Portela Fontoura, A.B., Javaid, A., de la Maza-Escol\u00e0, V.S., Salandy, N.S., Fubini, S.L., Grilli, E., McFadden, J.W., and Duan, J.E. (2023). Transcriptomic regulations of heat stress response in the liver of lactating dairy cows. BMC Genom., 24.","DOI":"10.1186\/s12864-023-09484-1"},{"key":"ref_144","doi-asserted-by":"crossref","unstructured":"Yue, S., Wang, Z., Wang, L., Peng, Q., and Xue, B. (2020). Transcriptome functional analysis of mammary gland of cows in heat stress and thermoneutral condition. Animals, 10.","DOI":"10.3390\/ani10061015"},{"key":"ref_145","doi-asserted-by":"crossref","unstructured":"Czech, B., Wang, Y., Wang, K., Luo, H., Hu, L., and Szyda, J. (2022). Host transcriptome and rumen microbiome interactions in Holstein cattle under heat stress. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.998093"},{"key":"ref_146","doi-asserted-by":"crossref","unstructured":"Zeng, H., Xia, H., Wang, X., Wang, Y., Fang, J., Li, S., Zhai, Y., and Han, Z. (2023). Comprehensive profiling of ceRNA (circRNA-miRNA-mRNA) networks in hypothalamic-pituitary-mammary gland axis of dairy cows under heat stress. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24010888"},{"key":"ref_147","doi-asserted-by":"crossref","unstructured":"Feng, X., Cheng, S., Lu, Z., Chen, X., Mu, T., Wang, C., Gu, Y., Li, Y., Chen, X., and Zheng, J. (2026). Construction of the ceRNA Regulatory Network Associated with Milk Fat Metabolism. Animals, 16.","DOI":"10.3390\/ani16040638"},{"key":"ref_148","doi-asserted-by":"crossref","unstructured":"Yu, X., Ceballos, D.A., Ferreira, M.M., Zapata, A., Seneviratne, N., Pokharel, S., Fang, Y., Li, G., and Leal-Yepes, F. (2026). Single-nucleus RNA sequencing reveals cell type-specific responses to heat stress in bovine mammary gland. bioRxiv.","DOI":"10.64898\/2026.02.18.706602"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"6637","DOI":"10.1038\/s41598-022-10461-z","article-title":"Carry-over effects of dry period heat stress on the mammary gland proteome and phosphoproteome in the subsequent lactation of dairy cows","volume":"12","author":"Skibiel","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"47","DOI":"10.5713\/ajas.16.0428","article-title":"Blood amino acids profile responding to heat stress in dairy cows","volume":"31","author":"Guo","year":"2018","journal-title":"Asian-Australas. J. Anim. Sci."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"101049","DOI":"10.1016\/j.animal.2023.101049","article-title":"Plasma and milk metabolomics reveal changes in amino acid metabolism in Holstein dairy cows under heat stress","volume":"18","author":"Leung","year":"2024","journal-title":"Animal"},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Yang, Y., Chen, Y., Hu, L., Zhang, C., Chen, G., Hou, L., Xu, Q., Wang, Y., and Li, M. (2025). Molecular regulation of whole genome DNA methylation in heat stress response of dairy cows. BMC Genom., 26.","DOI":"10.1186\/s12864-025-11683-x"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"11371","DOI":"10.1038\/s41598-021-89951-5","article-title":"Heat stress and immune response phenotype affect DNA methylation in blood mononuclear cells from Holstein dairy cows","volume":"11","author":"Livernois","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"1797","DOI":"10.1007\/s00484-022-02320-3","article-title":"Effect of acute heat shock on stress gene expression and DNA methylation in zebu (Bos indicus) and crossbred (Bos indicus \u00d7 Bos taurus) dairy cattle","volume":"66","author":"Masroor","year":"2022","journal-title":"Int. J. Biometeorol."},{"key":"ref_155","first-page":"6620","article-title":"Effects of micronutrient supplementation on performance and epigenetic status in dairy cows","volume":"105","author":"Gasselin","year":"2022","journal-title":"J. Dairy Sci."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"144550","DOI":"10.1016\/j.gene.2020.144550","article-title":"Effects of Chronic Heat Stress on mRNA and miRNA Expressions in Dairy Cows","volume":"742","author":"Liu","year":"2020","journal-title":"Gene"},{"key":"ref_157","doi-asserted-by":"crossref","unstructured":"Qi, Y., Zhang, Y., Zhang, J., Wang, J., and Li, Q. (2022). The alteration of N6-methyladenosine (m6A) modification at the transcriptome-wide level in response to heat stress in bovine mammary epithelial cells. BMC Genom., 23.","DOI":"10.1186\/s12864-022-09067-6"},{"key":"ref_158","first-page":"147","article-title":"Reducing water usage to cool cows by applying smart technologies","volume":"6","author":"Casarotto","year":"2025","journal-title":"JDS Commun."},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Du, D., Jiang, W., Feng, L., Zhang, Y., Chen, P., Wang, C., and Hu, Z. (2022). Effect of Saccharomyces cerevisiae culture in mitigating heat stress-related damage in dairy cows: A multi-omics characterisation. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.935004"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"501","DOI":"10.3168\/jds.2020-18653","article-title":"Predicting dairy cattle heat stress using machine learning techniques","volume":"104","author":"Becker","year":"2021","journal-title":"J. Dairy Sci."},{"key":"ref_161","doi-asserted-by":"crossref","unstructured":"Peretti, S., Rosa, V.D., Zotti, M., Prestes, A.M., Ferraz, P.F.P., da Silva, A.S., and Zotti, C.A. (2022). Thermoregulation and Performance of Dairy Cows Subjected to Different Evaporative Cooling Regimens, with or without Pepper Extract Supplementation. Animals, 12.","DOI":"10.3390\/ani12223180"},{"key":"ref_162","first-page":"186","article-title":"Effectiveness of cooling interventions on heat-stressed dairy cows based on a mechanistic thermoregulatory model","volume":"245","author":"Zhou","year":"2024","journal-title":"Biosyst. Eng."},{"key":"ref_163","doi-asserted-by":"crossref","unstructured":"Fournel, S., Ouellet, V., and Charbonneau, \u00c9. (2017). Practices for Alleviating Heat Stress of Dairy Cows in Humid Continental Climates: A Literature Review. Animals, 7.","DOI":"10.3390\/ani7050037"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"2687","DOI":"10.1007\/s11250-020-02309-3","article-title":"Effectiveness of Tunnel Ventilation as Dairy Cow Housing in Hot Climates: Rectal Temperatures during Heat Stress and Seasonal Variation in Milk Yield","volume":"52","author":"Dikmen","year":"2020","journal-title":"Trop. Anim. Health Prod."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"125","DOI":"10.3168\/jds.2009-2416","article-title":"The amount of shade influences the behavior and physiology of dairy cattle","volume":"93","author":"Rogers","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_166","unstructured":"Tucker, C.B., Drwencke, A., Van Os, J.M., Tresoldi, G., and Sch\u00fctz, K.E. (2019). Detecting and abating heat load in dairy cows. Proceedings of the Tri-State Dairy Nutrition Conference, Grand Wayne Center."},{"key":"ref_167","doi-asserted-by":"crossref","unstructured":"Moran, J.B. (2005). Tropical Dairy Farming: Feeding Management for Small Holder Dairy Farmers in the Humid Tropics, Landlinks Press.","DOI":"10.1071\/9780643093133"},{"key":"ref_168","doi-asserted-by":"crossref","unstructured":"Mylostyvyi, R., and Izhboldina, O. (2025). An integrated approach using temperature-humidity index, productivity, and welfare indicators for herd-level heat stress assessment in dairy cows. Animals, 15.","DOI":"10.3390\/ani15223341"},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"543","DOI":"10.2527\/jas1986.622543x","article-title":"Potential nutritional strategies for intensively managed cattle during thermal stress","volume":"62","author":"Beede","year":"1986","journal-title":"J. Anim. Sci."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"skac175","DOI":"10.1093\/jas\/skac175","article-title":"Increased supply of methionine during a heat-stress challenge in lactating Holstein cows alters mammary tissue mTOR signaling and its response to lipopolysaccharide","volume":"100","author":"Coleman","year":"2022","journal-title":"J. Anim. Sci."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"10851","DOI":"10.3168\/jds.2024-25225","article-title":"The Effects of Dietary Cation-Anion Difference and Dietary Buffer for Lactating Dairy Cattle under Mild Heat Stress with Night Cooling","volume":"107","author":"Bertens","year":"2024","journal-title":"J. Dairy Sci."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"4121","DOI":"10.3168\/jds.2009-2635","article-title":"Effect of saturated fatty acid supplementation on production and metabolism indices in heat-stressed mid-lactation dairy cows","volume":"93","author":"Wang","year":"2010","journal-title":"J. Dairy Sci."},{"key":"ref_173","doi-asserted-by":"crossref","unstructured":"Dunshea, F.R., Oluboyede, K., DiGiacomo, K., Leury, B.J., and Cottrell, J.J. (2019). Betaine improves milk yield in grazing dairy cows supplemented with concentrates at high temperatures. Animals, 9.","DOI":"10.3390\/ani9020057"},{"key":"ref_174","doi-asserted-by":"crossref","unstructured":"Williams, S.R.O., Milner, T.C., Garner, J.B., Moate, P.J., Jacobs, J.L., Hannah, M.C., Wales, W.J., and Marett, L.C. (2021). Dietary fat and betaine supplements offered to lactating cows affect dry matter intake, milk production, and body temperature responses to an acute heat challenge. Animals, 11.","DOI":"10.3390\/ani11113110"},{"key":"ref_175","doi-asserted-by":"crossref","unstructured":"Shah, A.M., Ma, J., Wang, Z., Zou, H., Hu, R., and Peng, Q. (2020). Betaine Supplementation Improves the Production Performance, Rumen Fermentation, and Antioxidant Profile of Dairy Cows in Heat Stress. Animals, 10.","DOI":"10.3390\/ani10040634"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"9745","DOI":"10.3168\/jds.2015-10514","article-title":"Evaluation of dietary betaine in lactating Holstein cows subjected to heat stress","volume":"99","author":"Hall","year":"2016","journal-title":"J. Dairy Sci."},{"key":"ref_177","first-page":"1657","article-title":"Effects of dietary betaine on body temperature indices, performance, metabolism, and hematological variables of dairy heifer calves during hot summer conditions","volume":"15","author":"Abuajamieh","year":"2022","journal-title":"Vet. World"},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1017\/S0007114521000908","article-title":"The effect of supplemental bioactive fatty acids on growth performance and immune function of milk-fed Holstein dairy calves during heat stress","volume":"127","author":"Mohtashami","year":"2022","journal-title":"Br. J. Nutr."},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1016\/j.livsci.2011.07.005","article-title":"Metabolic and hematological profiles in heat stressed lactating dairy cows fed diets supplemented with different selenium sources and doses","volume":"142","author":"Calamari","year":"2011","journal-title":"Livest. Sci."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"97","DOI":"10.21608\/jades.2016.67383","article-title":"Effect of selenium and vitamin E supplementation as a nutritional treatment for some physiological and productive traits of Holstein dairy cows under Egyptian summer conditions","volume":"17","author":"Khalifa","year":"2016","journal-title":"J. Egypt. Acad. Soc. Environ. Dev."},{"key":"ref_181","first-page":"2451","article-title":"Effects of microminerals on performance and metabolic adaptation in heat-stressed dairy cows: A review","volume":"203","author":"Lamanna","year":"2025","journal-title":"Biol. Trace Elem. Res."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1017\/S175173111200170X","article-title":"Nutrition, immune function and health of dairy cattle","volume":"7","author":"Ingvartsen","year":"2013","journal-title":"Animal"},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"114635","DOI":"10.1016\/j.anifeedsci.2020.114635","article-title":"Chromium Yeast Alleviates Heat Stress by Improving Antioxidant and Immune Function in Holstein Mid-Lactation Dairy Cows","volume":"269","author":"Shan","year":"2020","journal-title":"Anim. Feed Sci. Technol."},{"key":"ref_184","doi-asserted-by":"crossref","unstructured":"Chen, Y.-H., Chen, Y.-M., Tu, P.-A., Lee, K.-H., Chen, J.-Y., and Hsu, J.-T. (2023). Effect of Supplementing Vitamin E, Selenium, Copper, Zinc, and Manganese during the Transition Period on Dairy Cow Reproductive Performance and Immune Function. Vet. Sci., 10.","DOI":"10.3390\/vetsci10030225"},{"key":"ref_185","doi-asserted-by":"crossref","unstructured":"Son, A.-R., Kim, S.-H., Islam, M., Oh, S.-J., Paik, M.-J., Lee, S.-S., and Lee, S.-S. (2022). Higher Concentration of Dietary Selenium, Zinc, and Copper Complex Reduces Heat Stress-Associated Oxidative Stress and Metabolic Alteration in the Blood of Holstein and Jersey Steers. Animals, 12.","DOI":"10.3390\/ani12223104"},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"935","DOI":"10.3168\/jds.2008-1496","article-title":"Effects of a supplemental yeast culture on heat-stressed lactating Holstein cows","volume":"92","author":"Shwartz","year":"2009","journal-title":"J. Dairy Sci."}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/12\/918\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T01:22:52Z","timestamp":1781745772000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/12\/918"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,12]]},"references-count":186,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2026,6]]}},"alternative-id":["biology15120918"],"URL":"https:\/\/doi.org\/10.3390\/biology15120918","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,12]]}}}