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The thermoregulation system and how it works
Abstract
Heat exchange processes between the body and the environment are introduced. The definition of the thermoneutral zone as the ambient temperature range within which body temperature (T b) regulation is achieved only by nonevaporative processes is explained. Thermoreceptors, thermoregulatory effectors (both physiologic and behavioral), and neural pathways and T b signals that connect receptors and effectors into a thermoregulation system are reviewed. A classification of thermoeffectors is proposed. A consensus concept is presented, according to which the thermoregulation system is organized as a dynamic federation of independent thermoeffector loops. While the activity of each effector is driven by a unique combination of deep (core) and superficial (shell) T b s, the regulated variable of the system can be viewed as a spatially distributed T b with a heavily represented core and a lightly represented shell. Core T b is the main feedback; it is always negative. Shell T b s (mostly of the hairy skin) represent the auxiliary feedback, which can be negative or positive, and which decreases the system's response time and load error. Signals from the glabrous (nonhairy) skin about the temperature of objects in the environment serve as feedforward signals for various behaviors. Physiologic effectors do not use feedforward signals. The system interacts with other homeostatic systems by "meshing" with their loops. Coordination between different thermoeffectors is achieved through the common controlled variable, T b. The term balance point (not set point) is used for a regulated level of T b. The term interthreshold zone is used for a T b range in which no effectors are activated. Thermoregulatory states are classified, based on whether: T b is increased (hyperthermia) or decreased (hypothermia); the interthreshold zone is narrow (homeothermic type of regulation) or wide (poikilothermic type); and the balance point is increased (fever) or decreased (anapyrexia). During fever, thermoregulation can be either homeothermic or poikilothermic; anapyrexia is always a poikilothermic state. The biologic significance of poikilothermic states is discussed. As an example of practical applications of the concept presented, thermopharmacology is reviewed. Thermopharmacology uses drugs to modulate specific temperature signals at the level of a thermoreceptor (transient receptor potential channel).
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“While neurons responsive to cold temperatures also exist, they are less common and generally activated indirectly by warmth-sensitive neurons, emphasizing the greater risks associated with overheating as opposed to overcooling (Romanovsky, 2018).”
From Comparative analysis of energy transfer mechanisms for neural implants (2024) · cites this paper as Romanovsky, 2018
In the citing paper
“Adding another layer of complexity are specialized neurons in the preoptic anterior hypothalamus (POA), which are central to the brain’s thermal sensitivity.”
From Comparative analysis of energy transfer mechanisms for neural implants (2024) · cites this paper as Romanovsky, 2018
In the citing paper
“These neurons play a key role in physiological responses to temperature changes as most of these neurons are sensitive to warmth, increasing their activity as temperature rises triggering further regulations (Romanovsky, 2018).”
From Comparative analysis of energy transfer mechanisms for neural implants (2024) · cites this paper as Romanovsky, 2018
In the citing paper
“Moreover, some molecules, such as transient receptor potential (TRP) channels like TRPM2, are known to act as heat sensors in hypothalamic neurons (Romanovsky, 2018).”
From Comparative analysis of energy transfer mechanisms for neural implants (2024) · cites this paper as Romanovsky, 2018
In the citing paper
“While neurons responsive to cold temperatures also exist, they are less common and generally activated indirectly by warmth-sensitive neurons, emphasizing the greater risks associated with overheating as opposed to overcooling (Romanovsky, 2018).”
From Comparative analysis of energy transfer mechanisms for neural implants (2024) · cites this paper as Romanovsky, 2018
In the citing paper
“Despite a biphasic rhythm in core body temperature being such a well-established and recognized feature of the ovulatory menstrual cycle, the neural pathways that link the female reproductive and thermoregulatory systems remain incompletely understood 5.”
From Temperature regulation in women: Effects of the menstrual cycle (2020) · cites this paper as [5]
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Danker‐Hopfe, H., Bueno‐Lopez, A., Dorn, H., Schmid, G., Hirtl, R., & Eggert, T. (2020) · International Journal of Hygiene and Environmental Health · doi:10.1016/j.ijheh.2020.113550 · Cited by 17
Stereolithography 3D Printing of a Heat Exchanger for Advanced Temperature Control in Wire Myography
Kelava, L., Ivić, I. D., Pákai, E., Fekete, K., Maróti, P., Told, R., Ujfalusi, Z., & Garami, A. (2022) · Polymers · doi:10.3390/polym14030471 · Cited by 16
“, reptiles, birds and mammals) are euthermic and maintain their core body temperature in relatively narrow ranges by using behavioral and non-behavioral thermoregulation [30].”
BackgroundEnvironmental neurology in the tropics
Reis, J. J., Spencer, P. S., Román, G. C., & Buguet, A. (2020) · Journal of the Neurological Sciences · doi:10.1016/j.jns.2020.117287 · Cited by 14
Thermoregulatory reflex control of cutaneous vasodilation in healthy aging
Greaney, J. L., Stanhewicz, A. E., Wolf, S. T., & Kenney, W. L. (2020) · Temperature · doi:10.1080/23328940.2020.1832950 · Cited by 14
“In humans, the thermoregulatory system is governed by multiple interrelated independent neural reflex arcs [10].”
BackgroundCold-Water Immersion: Neurohormesis and Possible Implications for Clinical Neurosciences
López-Ojeda, W., & Hurley, R. A. (2024) · Journal of Neuropsychiatry · doi:10.1176/appi.neuropsych.20240053 · Cited by 14
McDonough, W., Rich, J., Aragon, I. V., Saleh, L. A., Boyd, A., Richter, A., Koloteva, A., & Richter, W. (2020) · Biochemical Pharmacology · doi:10.1016/j.bcp.2020.114158 · Cited by 14
McDonald, P., Brown, H. A., Topham, T. H., Kelly, M. K., Jardine, W. T., Carr, A. J., Sawka, M. N., Woodward, A. P., Clark, B., & Périard, J. D. (2025) · Comprehensive physiology · doi:10.1002/cph4.70017 · Cited by 13
Women have a greater cardiac vagal withdrawal to heat stress compared to men
Ferreira, F. C., Padilha, M. C. S. V., Tobadini, E., Bellocchi, C., Carandina, A., Montano, N., Soares, P. P. D. S., & Rodrigues, G. D. (2022) · Temperature · doi:10.1080/23328940.2022.2135354 · Cited by 15
“The core temperature control depends on a complex neural circuit of peripheral and central thermoreceptors, central integration (preoptic area/ anterior hypothalamus), and efferent responses to heat production and dissipation [12].”
BackgroundThe contribution of cutaneous thermal signals to bodily self-awareness
Salvato, G., Jenkinson, P. M., Sellitto, M., Crivelli, D., Crottini, F., Fazia, T., Squarza, S. A. C., Piano, M., Sessa, M., Gandola, M., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-55829-7 · Cited by 13
Freitag, L., Clijsen, R., Deflorin, C., Taube, W., Taeymans, J., & Hohenauer, E. (2021) · Frontiers in Sports and Active Living · doi:10.3389/fspor.2021.660092 · Cited by 12
“…be affected by the subcutaneous adipose tissue thickness, as well as environmental, hormonal, temporal, and nutritional factors and varies between different body parts (Jutte et al., 2001; Costello et al., 2012b; Garami and Székely, 2014; Adams et al., 2015; Romanovsky, 2018; Baker et al., 2020).”
BackgroundNamisnak, L. H., Khoshnevis, S., & Diller, K. R. (2022) · Journal of Biomechanical Engineering · doi:10.1115/1.4056110 · Cited by 10
Artificial homeostatic temperature regulation via bio-inspired feedback mechanisms
Feketa, P., Birkoben, T., Noll, M., Schaum, A., Meurer, T., & Kohlstedt, H. (2023) · Scientific Reports · doi:10.1038/s41598-023-31963-4 · Cited by 11
Fowler, R. C., Jeffries, O., Tallent, J., Theis, N., Heffernan, S. M., McNarry, M. A., Kilduff, L. P., & Waldron, M. (2020) · European Journal of Sport Science · doi:10.1080/17461391.2020.1739144 · Cited by 10
“These contrasting approaches provide markedly different afferent signals to the thermoregulatory centre, with the controlled water-suit and ventral forearm heating methods preventing typical dry cooling mechanisms and delivering a more focussed artificial stimulus, respectively (Romanovsky, 2018).”
BackgroundFerreira, F. C., Padilha, M. C. S. V., Tobadini, E., Carandina, A., Montano, N., Soares, P. P. D. S., & Rodrigues, G. D. (2021) · Physiological Measurement · doi:10.1088/1361-6579/ac1497 · Cited by 10
“…is dependent of a complex neural system in the preoptic area/anterior hypothalamus that integrates afferent information fromperipheral and central thermoreceptors, triggering efferent responses to control heat production and dissipation, and core temperature (Webb 1995, Romanovsky 2018).”
BackgroundHenkel, S., Frings‐Meuthen, P., Diekmann, C., Coenen, M., Stoffel‐Wagner, B., Németh, R., Pesta, D. H., & Egert, S. (2025) · Journal of Nutrition · doi:10.1016/j.tjnut.2025.01.013 · Cited by 9
“Vascular dilation increases heat dissipation to the surface, and blood circulates more rapidly from the warmer core to the outer periphery with an elevated HR, accelerating heat dissipation [34,35], which can also be a result of activation of the sympathetic nervous system [27].”
Pressnell, Z. S., Neilson, L. E., Tupone, D., Pfeiffer, R. F., & Safarpour, D. (2025) · Cells · doi:10.3390/cells14231910 · Cited by 9
“Conceptually, these POA neurons operate within a broader thermoregulatory architecture in which skin (shell) inputs provide rapid auxiliary feedback, while deep (core) temperature supplies the principal negative feedback that drives effector recruitment [7].”
InfluentialThe hyperthermic effect of central cholecystokinin is mediated by the cyclooxygenase-2 pathway
Kéringer, P., Füredi, N., Gaszner, B., Mikó, A., Pákai, E., Fekete, K., Olah, E., Kelava, L., Romanovsky, A. A., Rumbus, Z., et al. (2021) · American Journal of Physiology-Endocrinology and Metabolism · doi:10.1152/ajpendo.00223.2021 · Cited by 9
“The imprecise (poikilothermic) type of thermoregulation is further discussed elsewhere (43, 57).”
BackgroundSelf-citationColditz, I. G. (2021) · Frontiers in Animal Science · doi:10.3389/fanim.2021.652306 · Cited by 10
“Romanovsky (2018) describes the suite of effectors that influence a state or variable as constituting a dynamic confederation of independent controllers.”
BackgroundInfluentialSaleh, L. A., Boyd, A., Aragon, I. V., Koloteva, A., Spadafora, D., Mneimneh, W. S., Barrington, R. A., & Richter, W. (2021) · The FASEB Journal · doi:10.1096/fj.202100495r · Cited by 10
Water-filtered Infrared A (wIRA) Irradiation
Vaupel, P. W. (2022) · Springer eBooks · doi:10.1007/978-3-030-92880-3 · Cited by 10
de Andrade, M. T., Goulart, K. N. D. O., Barbosa, N. H. S., Soares, D. D., de Andrade, A. G. P., Gonçalves, D. A. P., Mendes, T. T., Coimbra, C. C., & Wanner, S. P. (2022) · Temperature · doi:10.1080/23328940.2022.2115274 · Cited by 9
“There are many definitions of the TNZ (for a detailed review, see ref [57]).”
BackgroundGhattassi, K., Farjallah, M. A., Graja, A., Romdhani, M., Boudhina, N., Guimard, A., Driss, T., Souissi, N., Chtourou, H., & Hammouda, O. (2024) · Research Quarterly for Exercise and Sport · doi:10.1080/02701367.2024.2303457 · Cited by 11
“MEL causes activation of central/peripheral vascular receptors which result in a vasodilation (Zhao et al., 2017) leading to a decrease in body temperature (Marrin et al., 2013; Romanovsky, 2018) which facilitates sleep onset (Szymusiak, 2018).”
Spontaneous and pharmacologically induced hypothermia protect mice against endotoxic shock
Tinajero, A., Merchant, W., Khan, A., Surbhi, Caron, A., Reynolds, R., Jia, L., & Gautron, L. (2025) · British Journal of Pharmacology · doi:10.1111/bph.70000 · Cited by 8
“…in laboratory rodents treated with a moderate dose of LPS (Corrigan et al., 2014; Ganeshan et al., 2019; Mei et al., 2018; Ogawa et al., 2016), here, we further establish that LPS-induced hypothermia during a lethal shock is a regulated and adaptive behavioural strategy with a survival benefit.”
InfluentialHomeostasis and Health: From Balance to Change
Bich, L., & Menatti, L. (2025) · Biological Theory · doi:10.1007/s13752-025-00510-x · Cited by 8
“…of mechanisms that measure deviations from a setpoint of certain variables such as temperature, glucose concentration, or body weight, has been deeply questioned in different branches of physiology (Müller et al. 2010; Speakman et al. 2011; Romanovsky 2018; Bich et this premodern family of ideas.”
The Role of Hyperthermia in the Treatment of Peritoneal Surface Malignancies
Guerra‐Londono, C. E., Tarazona, C. G., Sánchez-Monroy, J. A., Heppell, O., Guerra-Londoño, J. J., & Shah, R. A. (2022) · Current Oncology Reports · doi:10.1007/s11912-022-01275-3 · Cited by 7
Neural substrates underlying rhythmic coupling of female reproductive and thermoregulatory circuits
Grant, A. D., & Kriegsfeld, L. J. (2023) · Frontiers in Physiology · doi:10.3389/fphys.2023.1254287 · Cited by 7
“Although each part of the body comprises a “ micro-climate ” , BT is traditionally divided into the skin or “ shell ” , and the “ core ” or interior of the body (Childs, 2018; Romanovsky, 2018).”
BackgroundIncreasing intensity directly increases the perceived warmth of primary colors
Hammond, B. R., Gardner, C. R., Wooten, B. R., & Renzi‐Hammond, L. M. (2024) · Scientific Reports · doi:10.1038/s41598-024-77942-1 · Cited by 7
“Thermoregulation is metabolically expensive 23 .”
Langan, S. P., Casa, D. J., & Kwon, O. S. (2025) · Journal of Thermal Biology · doi:10.1016/j.jtherbio.2025.104331 · Cited by 6
Santiago, H. P., Leite, L. H. R., Lima, P. M. A., Fóscolo, D. R. C., Natali, A. J., Prímola‐Gomes, T. N., Szawka, R. E., & Coimbra, C. C. (2024) · Pflügers Archiv - European Journal of Physiology · doi:10.1007/s00424-024-02916-1 · Cited by 6
“On the efferent side, these mPOA cells tonically suppress skin vasoconstriction [30].”
InfluentialHigh Temperature Drives Topoisomerase Mediated Chromosomal Break Repair Pathway Choice
Ashour, M. E., Allam, W. R., Elsayed, W., Atteya, R., Elserafy, M., Magdeldin, S., Hassan, M. K., & El‐Khamisy, S. F. (2021) · Cancers · doi:10.3390/cancers13102315 · Cited by 6
A Conformable Two-Dimensional Resistance Temperature Detector for Measuring Average Skin Temperature
Namisnak, L. H., Khoshnevis, S., & Diller, K. R. (2021) · Journal of Medical Devices · doi:10.1115/1.4051442 · Cited by 6
Shakeel, C. S., Hassan, U., Ilyas, F., Zariwala, M. M., Ilyas, S. M., & Khan, S. J. (2021) · Proceedings of the Institution of Mechanical Engineers Part H Journal of Engineering in Medicine · doi:10.1177/09544119211051176 · Cited by 6
“The core body temperature of the human body is regulated via the hypothalamus.(4) It has been found that adequate clothing advancements and technology can aid in maintaining the optimum body temperature as well as the heart rate of the participating athlete.”
BackgroundDamasceno, J. R., Cavalcante, T. F., Ferreira, J. E. D. S. M., Barbosa, E. D. S., Moreira, R. P., Lopes, M. V. D. O., & Lira, A. L. B. D. C. (2022) · Revista Brasileira de Enfermagem · doi:10.1590/0034-7167-2021-0620 · Cited by 5
Medina, A., Méndez, J. I., Ponce, P., Peffer, T. E., Meier, A., & Molina, A. (2022) · Lecture notes in computer science · doi:10.1007/978-3-031-22061-6_18 · Cited by 5
Ishikawa, D. T., Vizin, R. C. L., de Souza, C. O., Carrettiero, D. C., Romanovsky, A. A., & Almeida, M. C. (2019) · Pharmaceuticals · doi:10.3390/ph12010024 · Cited by 5
“Both heat and cold defenses are represented by autonomic (or physiological; especially in endotherms) and behavioral (in all animals) effectors [2].”
BackgroundSelf-citationImpact of Reflective Roadways on Simulated Heat Strain at the Tokyo, Paris and Los Angeles Olympics
Vanos, J., Joshi, A., Guzmán-Echavarría, G., Rykaczewski, K., & Hosokawa, Y. (2024) · Journal of Science in Sport and Exercise · doi:10.1007/s42978-024-00294-9 · Cited by 5
“The ability to regulate body temperature depends on several factors such as age, underlying medical conditions, hydration, heat acclimation status, etc. [9, 16, 64, 69].”
Schaper, C. D. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/566679 · Cited by 5
“As described in the recent article heading a collection of research chapters on thermoregulation [24], theories of thermoregulation can tbe grouped into two categories: one historical”
BackgroundInfluentialCabanzo‐Olarte, L. C., Bícego, K. C., & Iannini, C. A. N. (2024) · Journal of Thermal Biology · doi:10.1016/j.jtherbio.2024.103889 · Cited by 4
Exercise performance effect of central dopamine is mediated by hypothalamic neuronal activation
Rodrigues, Q. T., Drummond, L. R., Lima, P. M. A., Machado, F. S. M., Campos, H. O., Szawka, R. E., Leite, L. H. R., & Coimbra, C. C. (2024) · Behavioural Brain Research · doi:10.1016/j.bbr.2024.115406 · Cited by 4
Metadata
- Authors
- Romanovsky, A. A.
- Affiliations
- St. Joseph's Hospital and Medical Center (US)
- Corresponding author
- Andrej A. Romanovsky
- ORCID
- Andrej A. Romanovsky
- Venue
- Handbook of Clinical Neurology
- Publisher
- Elsevier
- Published
- 2018
- Volume
- 156
- Pages
- 3-43
- ISSN
- 0072-9752 (ISSN-L)
- OpenAlex
- W2901310937
- Cited by
- 186
- References
- 220
- Citations by year
- 2026: 24 · 2025: 31 · 2024: 32 · 2023: 18 · 2022: 25 · 2021: 30 · 2020: 20 · 2019: 6
- FWCI
- 46.46
- Type
- book chapter
- Language
- en
- Open access
- closed
- Locations
- Handbook of clinical neurology (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Health Sciences › Medicine › Physiology (medical) › Thermoregulation and physiological responses
- Health Sciences › Medicine › Critical Care and Intensive Care Medicine › Thermal Regulation in Medicine
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Effects of Environmental Stressors on Livestock
- Keywords
- thermoregulation, thermoneutral zone, thermoreceptors, balance point, homeothermy, poikilothermy, fever, hyperthermia, hypothermia, TRP channels
- MeSH terms
- Animals, Body Temperature, Body Temperature Regulation, Humans, Neural Pathways, Feedback, Physiological
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