Abstract
Purpose
We examined whether short-term endurance training in the cold combined with layered clothing enhances cardiovascular, thermoregulatory, and performance adaptations.
Methods
Sixteen healthy males completed five consecutive days of endurance cycling in the cold (10 °C, 50% relative humidity [rH]) while wearing either thermal layered clothing (WEAR) or light clothing (CON). Before and after training, participants performed a graded exercise test, a time-to-exhaustion test, and a heat stress test (30 min at 35 °C, 50% rH).
Results
During training, heart rate, skin temperature, and sweat rate were higher in WEAR than CON (P < 0.05). Maximal power output increased similarly in both groups (WEAR: + 7.2 ± 9.5%; CON: + 8.4 ± 7.1%, P < 0.05), while maximal oxygen uptake improved only in WEAR (52.4 ± 7.3 to 55.2 ± 7.1 mL/kg/min, P < 0.05). During the heat stress test, ratings of perceived exertion and thermal sensation decreased after training in WEAR only (P < 0.05), with no changes in sweat rate, skin temperature, or rectal temperature in either group.
Conclusion
Short-term endurance training in the cold combined with layered clothing attenuated perceptual strain during subsequent exercise in the heat, implying a practical alternative approach to traditional heat acclimation strategies.
Keywords: Heat acclimation, Thermal strain, Thermoregulation, Clothing, Cycling, Endurance performance
Introduction
Exercising in hot environments impairs endurance performance and increases the risk of heat illness (Periard et al. 2021). To mitigate such adverse effects, athletes commonly undergo repeated heat exposure (i.e., heat acclimation) before competition in the heat. Traditional heat acclimation protocols typically involve 60–90 min of daily exercise in hot conditions over ~ 10–14 days (Periard et al. 2015). These interventions improve endurance capacity and heat tolerance through cardiovascular and thermoregulatory adaptations such as plasma volume expansion, increased sweat rate, and reductions in heart rate and core temperature (Periard et al. 2015, 2016).
Short-term (≤ 7 days) heat acclimation has gained increasing attention as a more practical alternative across many sports due to congested competition schedules and limited preparation time (Garrett et al. 2011; Chalmers et al. 2014). However, its implementation can be challenging in cold climates or during winter, particularly when environmental chambers are unavailable. To address this limitation, alternative approaches aimed at increasing thermal strain have been proposed, including sauna exposure, hot water immersion, and the use of thermal clothing (Heathcote et al. 2018; Stevens 2018; Jenkins et al. 2025). Among these methods, thermal clothing is particularly attractive because it requires no specialized facilities. For example, wearing an upper-body sauna suit increased peak rectal temperature (Trec) and sweat rate during 30 min of cycling under both temperate (22 °C) and hot (45 °C) conditions (Willmott et al. 2018). Moderate increases in core temperature and sweat rate were also observed during 80 min of outdoor cycling (~ 17 °C) when wearing thermal clothing compared with light clothing (Stevens et al. 2017). Likewise, treadmill running for 60 min in a cool (15 °C) environment while wearing multiple clothing layers elevated Trec above 38.5 ℃ (Ely et al. 2018), a threshold considered sufficient to induce heat acclimation adaptations (Fox et al. 1963).
In terms of training adaptations, an early review article highlighted that exercise training with thermal clothing is likely to improve heat tolerance, while evidence regarding effects beyond those induced by training itself is limited (Dawson 1994). Recently, Lundby et al. (2021) found that 10 days of cycling in a cool (18.4 °C) environment while wearing layered clothing significantly improved performance during a 30-min all-out cycling test in the heat, with effects comparable to training at 35℃ without additional clothing. Furthermore, 5 days of treadmill running in a temperate condition (20 °C) with overdressing induced greater reduction in resting and peak core temperature, skin temperature, and heart rate during 60-min running in the heat (40 °C) compared with the same training with light clothing (Greenfield et al. 2025). However, because these assessments were conducted under temperate conditions, it remains unclear whether comparable adaptations occur when training is undertaken in the cold (e.g., 10 °C). In cold conditions, a larger skin-environment thermal gradient enhances dry heat loss via convection and radiation, limiting core temperature rise (Blokker et al. 2022; Wallace et al. 2024). Wearing layered clothing restricts this heat dissipation, allowing metabolic heat production to elevate internal thermal strain. This combination may induce heat acclimation-like cardiovascular and thermoregulatory adaptations.
Therefore, this study tested the hypothesis that short-term endurance training in the cold (10 °C) while wearing layered clothing enhances cardiovascular, thermoregulatory, and performance adaptations compared with identical training performed in light clothing.
Methods
Participants
Sixteen recreationally active males (age: 20.8 ± 0.9 years, height: 172.7 ± 4.6 cm, body weight: 64.3 ± 7.2 kg) participated in the study. All participants were accustomed to low- to moderate-intensity cycling and had not performed any heat-based training for at least six months before the experiment. Participants received detailed information about the experimental procedures and provided written informed consent. The study was approved by the Research Ethics Committee of Ritsumeikan University, Japan (BKC-IRB-2020-048), and conducted in accordance with the Declaration of Helsinki.
Experimental overview
The study consisted of three performance tests – a graded exercise test (GXT), time-to-exhaustion (TTE) test, and heat stress test (HST) – conducted before and after five consecutive days of endurance training in the cold (Fig. 1). A five-day training regimen was chosen based on previous short-term heat acclimation programs as a practically applicable strategy for athletes (Garrett et al. 2011; Chalmers et al. 2014). Participants were randomly assigned to either a layered clothing group (WEAR, n = 8) or a light clothing group (CON, n = 8) for all training sessions. WEAR trained while wearing layered thermal clothing including caps, neck warmers, long-sleeved shirts, insulated jackets, long tights, insulated trousers, socks, and shoes. In contrast, CON wore standard (light clothing) attire including T-shirts, shorts, socks, and shoes.
Fig. 1.

Experimental overview of the short-term endurance training intervention performed with layered clothing (WEAR) or light clothing (CON), including pre- and post-training testing sequences. GXT graded exercise test, TTE time to exhaustion, HST heat stress test
Pre- and post-training tests were performed in a fixed order (GXT, TTE, HST) over three consecutive days, with two rest days following the final training session. Participants were instructed to refrain from caffeine, alcohol, or supplements and to avoid additional exercise from 24 h before the pre-training tests until completion of the post-training tests. All tests and training sessions were conducted in an environmental chamber (FCC-500S; Fuji Medical Science Co. Ltd., Chiba, Japan), with ambient temperature and relative humidity (rH) continuously monitored.
Pre- and post-training tests
Graded exercise test
The GXT was conducted on an electromagnetically braked cycle ergometer (Aerobike 75XLⅢ; Konami Corp., Tokyo, Japan) in a thermoneutral environment (25 °C, 50% rH) to determine maximal aerobic capacity. The test began at 70 W and increased by 30 W every 2 min until volitional exhaustion at a cadence of 70 rpm. Maximal power output (Wmax) was defined as the highest workload completed for ≥ 1 min. Expired gases were continuously collected and analyzed using an automatic gas analyzer (AE300S; Minato Medical Science Co. Ltd., Tokyo, Japan). Heart rate (HR), stroke volume (SV), and cardiac output (CO) were measured using an impedance cardiac output meter (PhysioFlow Q-Link; Manatec Biomedical, Paris, France). Six electrodes were placed on the neck, chest, and back. Cardiorespiratory data were averaged over 30-s intervals, and maximal values for oxygen uptake (VO2max), HR (HRmax), SV (SVmax), and CO (COmax) were determined.
Time-to-exhaustion test
In a thermoneutral environment (25 °C, 50% rH), participants performed a constant-load cycling test to volitional exhaustion (70 rpm) on a cycle ergometer (Aerobike 75XLⅢ; Konami Corp., Tokyo, Japan). Workload was fixed at 80% of VO2max (198 ± 38 W). The test was terminated when cadence dropped below 65 rpm for > 5 s, and time to exhaustion (TTE) was recorded.
Heat stress test
The HST consisted of 30 min of continuous cycling at 65% of VO2max (152 ± 31 W) in a hot environment (35 °C, 50% rH). HR was continuously monitored using a wireless HR monitor (RCX5; Polar Electro Oy., Kempele, Finland). Trec was measured using a thermistor (ITP010-11; Nikkiso Therm Co. Ltd., Tokyo, Japan) inserted 10 cm past the anal sphincter. Skin temperature was measured using surface thermistors (ITP082-24; Nikkiso Therm Co. Ltd.) attached to the left chest, arm, thigh, and calf. Mean skin temperature (Tskin) was calculated as Tskin = 0.3 × (chest + arm) + 0.2 × (thigh + calf) (Ramanathan 1964). Data were averaged over 5-min intervals. Ratings of perceived exertion (RPE, 10-point scale) and thermal sensation (TS, 9-point scale: 1 = very cold to 9 = very hot) were assessed every 10 min. Nude body weight was measured before and after the HST to estimate sweat rate.
Body composition and fluid balance
Body composition and fluid balance (including blood sampling) were assessed immediately before pre- and post-training GXT, following an overnight fast. Participants arrived at the laboratory at 8.30 am and rested for 10 min before measurements. Body weight, lean body mass, and body fat percentage were measured using a multi-frequency bioelectrical impedance analyzer (Inbody770; InBody Co. Ltd., Seoul, South Korea). Venous blood samples were collected using a 2.5 mL syringe, and hemoglobin (Hb) and hematocrit (Hct) concentrations were analyzed by a certified clinical laboratory (SRL Inc., Tokyo, Japan). Changes in plasma volume (ΔPV) were calculated as ΔPV = 100 × ([Hbpre/Hbpost] × [100 − Hctpost]/[100 − Hctpre] − 1) (Dill and Costill 1974). Urine samples were collected to assess urine specific gravity using a refractometer (PAL-09S; Atago Co. Ltd., Tokyo, Japan).
Training protocol
Participants completed five consecutive days of endurance cycling in a cold environment (10 °C, 50% rH). The ambient temperature of 10 °C represents the typical daily maximum temperature in winter in major cities in Japan (e.g., Tokyo, Osaka, and Nagoya). Each training session comprised four 20-min blocks, each including 5 min of cycling at 50 W followed by 15 min at 62% of Wmax, and ended with 20 min at 38% of Wmax (total duration of 100 min; Fig. 1). Pilot testing confirmed that this protocol elicited sufficient core temperature elevation (> 38.5 °C) when layered clothing was worn. During each 50 W stage of blocks 2–4, participants consumed 200 mL of water (20 °C; total 600 mL in both WEAR and CON) to minimize dehydration. HR, Trec, Tskin, sweat rate, RPE, and TS were monitored on training days 1, 3, and 5. The cumulative heat strain index (CHSI) was calculated from HR and Trec during each training session as follows (Frank et al. 2001):
Where: hb = heart beats, fc0 = baseline HR (bpm), Trec0 = baseline Trec ( °C), and t = time (min) from the onset of exercise.
Statistical analyses
Data are expressed as mean ± standard deviation (SD). Two-way repeated-measures ANOVA (group × period [pre- and post-training]) was used to compare body composition and fluid balance, performance and physiological variables during the GXT, TTE, and training sessions (SPSS; IBM Corp., Armonk, USA). Physiological and perceptual responses during the HST were analyzed using two-way repeated-measures ANOVA (period [pre- and post-training] × time [baseline to 30 min]). Effect sizes were calculated as partial eta squared (ηp2), and values of 0.01, 0.06, and > 0.14 were considered as small, medium, and large, respectively (Cohen 1988). When significant main effects or interaction were observed, Tukey post hoc tests were performed. Statistical significance was set at P < 0.05.
Results
Training data
Table 1,Summarizes physiological and perceptual responses during the training sessions. Peak and mean HR were significantly higher in WEAR than CON (main effect of group: P < 0.001 [ηp2 = 0.643] for peak HR; P = 0.001 [ηp2 = 0.567] for mean HR), with significant reductions from day 1 to day 5 observed only in WEAR (main effect of period: P < 0.001 [ηp2 = 0.475] for peak HR; P < 0.001 [ηp2 = 0.474] for mean HR). Peak and mean Tskin were also higher in WEAR than in CON throughout the training period (main effect of group: P < 0.001 [ηp2 = 0.838] for peak Tskin; P < 0.001 [ηp2 = 0.893] for mean Tskin). Peak Trec was significantly higher in WEAR on day 1, but this difference diminished on subsequent days (main effect of group: P = 0.017 [ηp2 = 0.363]). Within WEAR, both peak and mean Trec decreased from day 1 to day 5, whereas no changes were observed in CON (main effect of period: P = 0.002 [ηp2 = 0.370] for peak Trec; P = 0.023 [ηp2 = 0.253] for mean Trec). CHSI was significantly higher in WEAR than in CON on day 1 and 3, but not on day 5 (main effect of group: P = 0.008 [ηp2 = 0.407]). Sweat rate, RPE, and TS were consistently higher in WEAR than in CON (main effect of group: P = 0.007 [ηp2 = 0.415], P = 0.003 [ηp2 = 0.481], and P < 0.001 [0.665], respectively).
Table 1.
Physiological and perceptual responses during training sessions in the layered clothing (WEAR) and light clothing (CON) groups
| WEAR group | CON group | |||||
|---|---|---|---|---|---|---|
| Day 1 | Day 3 | Day 5 | Day 1 | Day 3 | Day 5 | |
| HR (bpm) | ||||||
| Peak | 181 ± 7† | 174 ± 7*† | 171 ± 9*† | 161 ± 10 | 156 ± 9 | 155 ± 5 |
| Mean | 159 ± 9† | 154 ± 7† | 150 ± 8*† | 140 ± 9 | 137 ± 9 | 136 ± 6 |
| Trec (℃) | ||||||
| Peak | 38.9 ± 0.7† | 38.6 ± 0.4 | 38.4 ± 0.4* | 38.2 ± 0.2 | 38.1 ± 0.2 | 38.0 ± 0.2 |
| Mean | 38.2 ± 0.5 | 38.2 ± 0.3 | 37.9 ± 0.3* | 37.9 ± 0.2 | 37.8 ± 0.2 | 37.8 ± 0.2 |
| Tskin (℃) | ||||||
| Peak | 37.1 ± 0.9† | 36.8 ± 0.6† | 36.8 ± 0.4† | 34.2 ± 1.4 | 34.1 ± 0.9 | 34.3 ± 0.9 |
| Mean | 36.1 ± 1.0† | 35.9 ± 0.7† | 35.9 ± 0.4† | 31.9 ± 1.9 | 31.6 ± 1.6 | 31.7 ± 2.3 |
| CHSI (units) | 428 ± 60† | 315 ± 34*† | 296 ± 39* | 252 ± 40 | 184 ± 49 | 215 ± 59 |
| Sweat rate (L/h) | 1.0 ± 0.4† | 1.1 ± 0.5† | 1.1 ± 0.3† | 0.6 ± 0.2 | 0.6 ± 0.2 | 0.6 ± 0.2 |
| RPE | 7.5 ± 2.1† | 7.0 ± 1.7† | 6.9 ± 1.5† | 4.6 ± 1.3 | 4.6 ± 1.6 | 4.3 ± 1.6 |
| TS | 7.5 ± 1.8† | 7.5 ± 1.9† | 7.5 ± 1.4† | 3.1 ± 2.1 | 3.1 ± 2.0 | 3.1 ± 1.5 |
HR heart rate, Trec rectal temperature, Tskin mean skin temperature, CHSI cumulative heat strain index, RPE rating of perceived exertion, TS thermal sensation, Values are mean ± SD
*P < 0.05 vs. Day 1
†P < 0.05 vs. CON group
Body composition and fluid balance
No significant pre- to post-training changes were noted in either group for body weight (64.3 ± 9.8 to 64.2 ± 9.8 kg in WEAR; 64.2 ± 4.1 to 63.9 ± 4.2 kg in CON), lean body mass (55.5 ± 8.1 to 55.1 ± 8.1 kg in WEAR; 56.4 ± 4.3 to 55.7 ± 4.0 kg in CON), body fat percentage (13.7 ± 3.8 to 14.0 ± 3.9% in WEAR; 12.1 ± 2.3 to 12.8 ± 2.3% in CON), urine specific gravity (1.022 ± 0.009 to 1.024 ± 0.009 in WEAR; 1.022 ± 0.010 to 1.018 ± 0.008 in CON), Hb (15.2 ± 0.6 to 15.3 ± 0.8 g/dL in WEAR; 15.0 ± 0.9 to 15.2 ± 0.8 g/dL in CON), Hct (46.0 ± 1.4 to 46.2 ± 2.6% in WEAR; 45.5 ± 2.4 to 46.0 ± 1.9% in CON), or ΔPV (−0.7 ± 9.0% in WEAR; −2.0 ± 4.8% in CON).
Graded exercise test
As shown in Table 2, Wmax increased from pre- to post-training in both groups (+ 7.2 ± 9.5% and + 8.4 ± 7.1% in WEAR and CON, respectively; main effect of period: P = 0.002 [ηp2 = 0.521]). Absolute (+ 5.3 ± 4.6%) and relative VO2max (+ 5.6 ± 3.8%) increased in WEAR only (main effect of period: P = 0.009 [ηp2 = 0.400] and P = 0.004 [ηp2 = 0.464], respectively). HRmax (−1.4 ± 2.5% in WEAR and −1.8 ± 4.3% in CON), SVmax (+ 13.2 ± 21.2% in WEAR and −2.8 ± 12.9% in CON), and COmax (+ 11.6 ± 19.7% in WEAR and −1.8 ± 15.5% in CON) did not differ significantly within or between groups.
Table 2.
Performance and cardiorespiratory responses during the graded exercise test in the layered clothing (WEAR) and light clothing (CON) groups
| WEAR group | CON group | ||||
|---|---|---|---|---|---|
| Pre | Post | Pre | Post | ||
| Wmax (W) | 269 ± 36 | 288 ± 42* | 258 ± 59 | 276 ± 49* | |
| Absolute VO2max (mL/min) | 3365 ± 510 | 3544 ± 568* | 3253 ± 621 | 3344 ± 523 | |
| Relative VO2max (mL/kg/min) | 52.4 ± 7.3 | 55.2 ± 7.1* | 50.8 ± 7.8 | 52.0 ± 5.8 | |
| HRmax (bpm) | 197 ± 9 | 194 ± 11 | 189 ± 10 | 187 ± 9 | |
| SVmax (mL) | 106 ± 18 | 118 ± 21 | 122 ± 23 | 121 ± 27 | |
| COmax (L/min) | 20.1 ± 2.8 | 22.1 ± 3.2 | 22.4 ± 4.1 | 21.7 ± 4.5 | |
Wmax maximal power output, VO2max maximal oxygen uptake, HRmax maximal heart rate, SVmax maximal stroke volume, COmax maximal cardiac output
Values are mean ± SD
*P < 0.05 vs. Pre
Time-to-exhaustion test
The TTE improved significantly after training in both groups (from 21.5 ± 9.2 to 30.7 ± 14.1 min [+ 46.5 ± 65.6%] in WEAR and from 25.4 ± 8.0 to 37.8 ± 14.4 min [+ 51.4 ± 49.1%] in CON; main effect of period: P = 0.001 [ηp2 = 0.573]), with no between-group difference.
Heat stress test
Peak HR decreased significantly after training in both groups (WEAR: −5.9 ± 4.8%, main effect of period: P = 0.032 [ηp2 = 0.563]; CON: −4.3 ± 2.9%, main effect of period: P = 0.008 [ηp2 = 0.662]). Peak Trec (WEAR: −0.3 ± 0.7%; CON: −0.3 ± 0.9%) and Tskin (WEAR: + 1.4 ± 3.4%; CON: −0.4 ± 1.5%) did not change (Fig. 2). RPE and TS scores decreased significantly after training in WEAR only (main effect of period: P = 0.013 [ηp2 = 0.613] and P = 0.007 [ηp2 = 0.667], respectively; Fig. 3). Sweat rate was unchanged in both groups (+ 15.1 ± 27.4% and + 4.2 ± 24.1% in WEAR and CON, respectively; main effect of period: P = 0.234 [ηp2 = 0.100]).
Fig. 2.

Heart rate (A), rectal temperature (B), and mean skin temperature (C) during the heat stress test in the layered clothing (WEAR) and light clothing (CON) groups. Values are mean ± SD. †P < 0.05 vs. post
Fig. 3.

Rating of perceived exertion (A) and thermal sensation (B) during the heat stress test in the layered clothing (WEAR) and light clothing (CON) groups. Values are mean ± SD. †P < 0.05 vs. post
Discussion
We examined the effects of wearing layered clothing during five consecutive days of endurance training in a cold environment. The main findings were that this strategy improved VO2max, did not affect TTE in a thermoneutral environment, and attenuated perceptual strain during a HST without concomitant changes in thermoregulatory responses.
Elevating core temperature above 38.5°C is considered a prerequisite for effective heat acclimation (Fox et al. 1963). During the first training session, peak Trec in WEAR exceeded this threshold and remained elevated for a substantial duration despite the cold environment (10°C), reaching levels comparable to those observed during traditional heat acclimation interventions (Lorenzo et al. 2010; Moss et al. 2020) and exercise while wearing thermal clothing in temperate environments (Ely et al. 2018; Lundby et al. 2021). These findings indicate that the present training strategy successfully induced meaningful heat strain during early exposure. However, peak Trec gradually decreased across sessions and no longer reached 38.5°C by the final training day. Although this reduction is consistent with the onset of acclimation, the reduced heat stimulus in later sessions may explain the absence of thermoregulatory adaptations (Trec, Tskin, sweat rate) during the HST in WEAR. Stevens et al. (2018) examined performance and physiological adaptations following 2 weeks of field-based training (six running/cycling sessions per week) in a temperate environment (~ 18 °C, ~ 67% rH) while wearing light clothing or thermal clothing. However, they found no benefit of wearing thermal clothing on 40-min time trial performance or thermoregulatory adaptation in the heat (32.5°C, 55% rH). Although absolute core temperature data during training sessions were not reported, they suggested that thermophysiological strain was insufficient to elicit heat acclimation, as mean HR during training was similar between groups (139 bpm). In contrast, Greenfield et al. (2025) conducted 5 days of treadmill-based training in a temperate condition (20°C) while participants wore either thermal or light clothing, and found higher peak Trec, Tskin, HR, and sweat rate with thermal clothing than light clothing during both the initial and final training sessions. Notably, peak Trec remained elevated to 38.5°C with thermal clothing even during the final session (Greenfield et al. 2025). Furthermore, the 5-day training with thermal clothing induced significant thermoregulatory and cardiorespiratory adaptations (i.e., reductions of resting and peak core temperature, skin temperature, and heart rate) during 60 min of treadmill running in the heat (40°C) (Greenfield et al. 2025). Our results showed markedly higher HR, Tskin, and CHSI in WEAR than CON over the training period; however, the difference in Trec diminished during the latter training sessions. Maintaining sufficient heat stress throughout training –via higher exercise intensity or greater insulation –may therefore be required to elicit robust thermoregulatory adaptations.
In contrast to the unchanged thermoregulatory responses, perceptual strain (RPE and TS) was reduced after five consecutive days of training. Although Tskin is closely associated with TS and thermal comfort (Song et al. 2025), the perceptual improvements observed here were not accompanied by changes in Trec or Tskin. This dissociation aligns with previous findings showing continued reductions in TS and improvements in thermal comfort when heat acclimation was extended from 5 to 10 days, despite plateaued Trec and Tskin after five days (Moss et al. 2020), implying that perceptual adaptations may occur independently of thermoregulatory adjustments. Lower RPE and TS at a given workload decrease subjective strain, whereas elevated effort and thermal perception impair self-paced performance in the heat (Flouris and Schlader 2015). Moreover, exacerbated thermal sensation has been shown to impair cognitive performance during exercise in the heat (Qin et al. 2025). Although exercise capacity during the HST was not directly assessed, short-term cold training with layered clothing may nonetheless enhance endurance performance in hot environments by attenuating perceptual strain compared to training with light clothing.
Another notable finding is that VO2max increased significantly after training in WEAR (+ 5.6 ± 3.8%) but not in CON (+ 3.0 ± 6.1%). Traditional 10-day heat acclimation protocols improve VO2max and COmax, and these effects are largely attributed to plasma volume expansion (Lorenzo et al. 2010), although improvements in VO2max remains inconsistent across studies (Karlsen et al. 2015; Keiser et al. 2015). In our study, plasma volume did not increase, likely because the heat strain induced by layered clothing was insufficient to evoke this response. Nevertheless, the magnitude of increases in VO2max (+ 5.6%) and COmax (+ 2.0 L/min) in WEAR was comparable to values reported following traditional heat acclimation (Lorenzo et al. 2010). The contribution of plasma volume expansion in mediating VO2max improvements remains debated, as increases in plasma volume without concurrent increases in hemoglobin mass may cause hemodilution without enhancing oxygen delivery (Warburton et al. 2000). Alternative mechanisms may therefore explain the observed VO2max increase, including augmented muscle vasodilatory capacity (Richey et al. 2022), optimized blood flow redistribution (Chen et al. 2013), and enhanced muscle oxidative capacity (Patton et al. 2018). These adaptations could facilitate oxygen delivery and utilization during maximal exercise, partially accounting for the improved aerobic capacity. Higher HR through training sessions in WEAR represents augmented physiological strain (internal training load), which may support cardiovascular adjustments, as the time spent under high internal load is considered crucial to improve VO2max (Buchheit and Laursen 2013). Consistent with this interpretation, Lundby et al. (2021) also reported improved aerobic capacity after endurance training with thermal clothing in the absence of hematological and thermoregulatory adaptations. Further research is needed to elucidate the mechanisms underpinning these responses.
Both Wmax during the GXT and TTE at 80% VO2max significantly improved after training in WEAR and CON, indicating that the training stimulus itself was sufficient to enhance performance, particularly given that participants were not accustomed to high-intensity cycling. These general training adaptations may have masked any additional effects of layered clothing, and the observed improvement in VO2max may be partially attributable to exercise training itself. Therefore, caution is needed when interpreting these results. Importantly, exercise capacity (i.e., Wmax and TTE) was assessed only under thermoneutral conditions. Meta-analytic evidence indicates that the performance benefits of heat acclimation are small when aerobic capacity is measured in thermoneutral conditions, but moderate to large when measured in the heat (Waldron et al. 2021). Thus, training with layered clothing may confer greater improvements in exercise capacity measured under hot conditions compared with light clothing.
The present study has several limitations. First, only male participants were recruited. A recent review highlighted that the magnitude of physiological adaptations (e.g., HR and core temperature) following heat acclimation is comparable between males and females (Wickham et al. 2021), and females can adapt to heat regardless of menstrual cycle phase (Lundby et al. 2023; Richard et al. 2025). On the other hand, females may require a longer heat acclimation period to achieve physiological adaptations comparable to those observed in males (Wickham et al. 2021), suggesting that a 5-day training program may be insufficient in females. In addition, the menstrual cycle influences both resting and exercising core temperatures in the heat (Christison et al. 2025), which could complicate the interpretations of training-induced changes in core temperature. For these reasons, only male participants were included in the present study. However, the next step should be to examine the effects of short-term endurance training with layered clothing in females. Second, the sample size was relatively small (n = 8 per group) because all experiments were completed within a single season to minimize confounding effects of natural heat acclimation. Third, the training duration was short (5 days), which may be insufficient compared to traditional heat-acclimation protocols (~ 10–14 days). Fourth, post-tests were performed after a two-day rest period following completion of training to minimize the influence of fatigue resulting from relatively high exercise intensity compared with typical heat acclimation protocols, as well as differences in internal load between groups. However, several physiological adaptations may be more pronounced immediately after training and may therefore have been underestimated in the present study (Daanen et al. 2018). Finally, exercise capacity was assessed only under thermoneutral conditions; future studies should evaluate exercise capacity in differing hot environments (i.e., hot-dry and hot-humid) to enhance practical relevance.
In conclusion, short-term endurance training with layered clothing improved VO2max and attenuated perceptual strain during exercise in the heat, without inducing measurable cardiovascular or thermoregulatory adaptations. This strategy may offer a practical and time-efficient preparatory strategy for athletes training for competition in hot conditions.
Acknowledgements
We appreciate all the participants who completed the experiments.
Abbreviations
- ANOVA
Analysis of variance
- CO
Cardiac output
- COmax
Maximal cardiac output
- CON
Light clothing group
- CHSI
Cumulative heat strain index
- GXT
Graded exercise test
- HR
Heart rate
- HRmax
Maximal heart rate
- HST
Heat stress test
- Hb
Hemoglobin
- Hct
Hematocrit
- RPE
Ratings of perceived exertion
- SD
Standard deviation
- SV
Stroke volume
- SVmax
Maximal stroke volume
- TS
Thermal sensation
- TTE
Time to exhaustion
- Trec
Rectal temperature
- Tskin
Mean skin temperature
- VO2max
Maximal oxygen uptake
- WEAR
Layered clothing group
- Wmax
Maximal power output
- rH
Relative humidity
- ΔPV
Plasma volume change
- ηp2
Partial eta squared
Author contributions
KY, AM, RY, and KG designed the research. KY, SO, and AM conducted experiments and analyses. KY wrote the manuscript, and OG and KG revised it. All authors approved the final manuscript.
Funding
Open Access funding provided by Ritsumeikan University. The present study was funded by Panasonic Corporation.
Declarations
Conflict of interest
RY is an employee of Panasonic Corporation. All other authors declare no conflicts of interest.
Footnotes
Publisher’s Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
References
- Blokker T, Bucher E, Steiner T, Wehrlin JP (2022) Effect of cold ambient temperature on heat flux, skin temperature, and thermal sensation at different body parts in elite biathletes. Front Sports Act Living 4:966203. 10.3389/fspor.2022.966203 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Buchheit M, Laursen PB (2013) High-intensity interval training, solutions to the programming puzzle: Part I: cardiopulmonary emphasis. Sports Med 43(5):313–338. 10.1007/s40279-013-0029-x [DOI] [PubMed] [Google Scholar]
- Chalmers S, Esterman A, Eston R, Bowering KJ, Norton K (2014) Short-term heat acclimation training improves physical performance: a systematic review, and exploration of physiological adaptations and application for team sports. Sports Med 44(7):971–988. 10.1007/s40279-014-0178-6 [DOI] [PubMed] [Google Scholar]
- Chen TI, Tsai PH, Lin JH, Lee NY, Liang MT (2013) Effect of short-term heat acclimation on endurance time and skin blood flow in trained athletes. Open Access J Sports Med 4:161–170. 10.2147/OAJSM.S45024 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Christison KS, Gurney SC, Williamson-Reisdorph CM, Covington AC, Dumke CL (2025) Menstrual cycle effects on thermoregulation while exercising in the heat. J Therm Biol 127:104036. 10.1016/j.jtherbio.2024.104036 [DOI] [PubMed] [Google Scholar]
- Cohen J (1988) Statistical power analysis for the behavioral sciences, Second Edition. Lawrence Erlbaum Associates, Hillsdale (NJ). 10.4324/9780203771587 [DOI] [Google Scholar]
- Daanen HAM, Racinais S, Periard JD (2018) Heat Acclimation Decay and Re-Induction: A Systematic Review and Meta-Analysis. Sports Med 48(2):409–430. 10.1007/s40279-017-0808-x [DOI] [PMC free article] [PubMed] [Google Scholar]
- Dawson B (1994) Exercise training in sweat clothing in cool conditions to improve heat tolerance. Sports Med 17(4):233–244. 10.2165/00007256-199417040-00004 [DOI] [PubMed] [Google Scholar]
- Dill DB, Costill DL (1974) Calculation of percentage changes in volumes of blood, plasma, and red cells in dehydration. J Appl Physiol 37(2):247–248. 10.1152/jappl.1974.37.2.247 [DOI] [PubMed] [Google Scholar]
- Ely BR, Blanchard LA, Steele JR, Francisco MA, Cheuvront SN, Minson CT (2018) Physiological responses to overdressing and exercise-heat stress in trained Runners. Med Sci Sports Exerc 50(6):1285–1296. 10.1249/MSS.0000000000001550 [DOI] [PubMed] [Google Scholar]
- Flouris AD, Schlader ZJ (2015) Human behavioral thermoregulation during exercise in the heat. Scand J Med Sci Sports 25(Suppl 1):52–64. 10.1111/sms.12349 [DOI] [PubMed] [Google Scholar]
- Fox RH, Goldsmith R, Kidd DJ, Lewis HE (1963) Acclimatization to heat in man by controlled elevation of body temperature. J Physiol 166(3):530–547. 10.1113/jphysiol.1963.sp007121 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Frank A, Belokopytov M, Shapiro Y, Epstein Y (2001) The cumulative heat strain index–a novel approach to assess the physiological strain induced by exercise-heat stress. Eur J Appl Physiol 84(6):527–532. 10.1007/s004210000368 [DOI] [PubMed] [Google Scholar]
- Garrett AT, Rehrer NJ, Patterson MJ (2011) Induction and decay of short-term heat acclimation in moderately and highly trained athletes. Sports Med 41(9):757–771. 10.2165/11587320-000000000-00000 [DOI] [PubMed] [Google Scholar]
- Greenfield AM, Brazelton SC, Alba BK, Bodurtha PO, Bradbury KE, Caldwell AR, Chapman CL, Charkoudian N, Connor KR, Conz K, Fry BA, Giersch GEW, Gonzalez-Rojas DH, Heikkinen ME, Looney DP, Mayer TA, McCarthy KG, Potter AW, Rioux TP, Salgado RM, Seeley AD, Shaw MA, Xu X, Ryan BJ (2025) Exercise with overdressing for heat acclimation: a multilayered approach using biophysical modeling and two randomized crossover trials. J Appl Physiol 139(4):889–901. 10.1152/japplphysiol.00624.2025 [DOI] [PubMed] [Google Scholar]
- Heathcote SL, Hassmen P, Zhou S, Stevens CJ (2018) Passive heating: reviewing practical heat acclimation strategies for endurance athletes. Front Physiol 9:1851. 10.3389/fphys.2018.01851 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jenkins EJ, Killick JA, Zerilli O, Douglas AJM, Corr L, Hughes MG, Tremblay JC, Stembridge M (2025) Long-term passive heat acclimation enhances maximal oxygen consumption via haematological and cardiac adaptation in endurance runners. J Physiol. 10.1113/JP289874 [DOI] [PubMed] [Google Scholar]
- Karlsen A, Racinais S, Jensen MV, Norgaard SJ, Bonne T, Nybo L (2015) Heat acclimatization does not improve VO2max or cycling performance in a cool climate in trained cyclists. Scand J Med Sci Sports 25(Suppl 1):269–276. 10.1111/sms.12409 [DOI] [PubMed] [Google Scholar]
- Keiser S, Fluck D, Huppin F, Stravs A, Hilty MP, Lundby C (2015) Heat training increases exercise capacity in hot but not in temperate conditions: a mechanistic counter-balanced cross-over study. Am J Physiol Heart Circ Physiol 309(5):H750-761. 10.1152/ajpheart.00138.2015 [DOI] [PubMed] [Google Scholar]
- Lorenzo S, Halliwill JR, Sawka MN, Minson CT (2010) Heat acclimation improves exercise performance. J Appl Physiol 109(4):1140–1147. 10.1152/japplphysiol.00495.2010 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lundby C, Svendsen IS, Urianstad T, Hansen J, Ronnestad BR (2021) Training wearing thermal clothing and training in hot ambient conditions are equally effective methods of heat acclimation. J Sci Med Sport 24(8):763–767. 10.1016/j.jsams.2021.06.005 [DOI] [PubMed] [Google Scholar]
- Lundby C, Hamarsland H, Hansen J, Bjorndal H, Berge SN, Hammarstom D, Ronnestad BR (2023) Hematological, skeletal muscle fiber, and exercise performance adaptations to heat training in elite female and male cyclists. J Appl Physiol 135(1):217–226. 10.1152/japplphysiol.00115.2023 [DOI] [PubMed] [Google Scholar]
- Moss JN, Bayne FM, Castelli F, Naughton MR, Reeve TC, Trangmar SJ, Mackenzie RWA, Tyler CJ (2020) Short-term isothermic heat acclimation elicits beneficial adaptations but medium-term elicits a more complete adaptation. Eur J Appl Physiol 120(1):243–254. 10.1007/s00421-019-04269-5 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Patton MG, Gillum TL, Szymanski MC, Gould LM, Lauterbach CJ, Vaughan RA, Kuennen MR (2018) Heat acclimation increases mitochondrial respiration capacity of C2C12 myotubes and protects against LPS-mediated energy deficit. Cell Stress Chaperones 23(5):871–883. 10.1007/s12192-018-0894-1 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Periard JD, Racinais S, Sawka MN (2015) Adaptations and mechanisms of human heat acclimation: applications for competitive athletes and sports. Scand J Med Sci Sports 25(Suppl 1):20–38. 10.1111/sms.12408 [DOI] [PubMed] [Google Scholar]
- Periard JD, Travers GJS, Racinais S, Sawka MN (2016) Cardiovascular adaptations supporting human exercise-heat acclimation. Auton Neurosci 196:52–62. 10.1016/j.autneu.2016.02.002 [DOI] [PubMed] [Google Scholar]
- Periard JD, Eijsvogels TMH, Daanen HAM (2021) Exercise under heat stress: thermoregulation, hydration, performance implications, and mitigation strategies. Physiol Rev 101(4):1873–1979. 10.1152/physrev.00038.2020 [DOI] [PubMed] [Google Scholar]
- Qin F, Hu K, Wu Y, Zheng X (2025) Single and combined effects of environmental heat stress and physical exercise on thermoregulation, executive function, and cerebral oxygenation. J Therm Biol 127:104057. 10.1016/j.jtherbio.2025.104057 [DOI] [PubMed] [Google Scholar]
- Ramanathan NL (1964) A new weighting system for mean surface temperature of the human body. J Appl Physiol 19:531–533. 10.1152/jappl.1964.19.3.531 [DOI] [PubMed] [Google Scholar]
- Richard NA, Cheung SS, Koehle MS, Claydon VE, Fenuta AM, Cote AT (2025) Three weeks of heat maintenance potentiates the benefits of heat acclimation in trained females. Physiol Rep 13(22):e70631. 10.14814/phy2.70631 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Richey RE, Hemingway HW, Moore AM, Olivencia-Yurvati AH, Romero SA (2022) Acute heat exposure improves microvascular function in skeletal muscle of aged adults. Am J Physiol Heart Circ Physiol 322(3):H386–H393. 10.1152/ajpheart.00645.2021 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Song W, Zhong F, Calautit JK, Li J (2025) Exploring the role of skin temperature in thermal sensation and thermal comfort: A comprehensive review. Energy and Built Environment 6(4):762–781. 10.1016/j.enbenv.2024.03.002 [DOI] [Google Scholar]
- Stevens CJ (2018) Evidence for training in additional clothing as an alternative heat acclimation strategy for athletes. Temperature (Austin) 5(2):101–102. 10.1080/23328940.2018.1437309 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stevens CJ, Plews DJ, Laursen PB, Kittel AB, Taylor L (2017) Acute physiological and perceptual responses to wearing additional clothing while cycling outdoors in a temperate environment:A practical method to increase the heat load. Temperature (Austin) 4(4):414–419. 10.1080/23328940.2017.1365108 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stevens CJ, Heathcote SL, Plews DJ, Laursen PB, Taylor L (2018) Effect of two-weeks endurance training wearing additionalclothing in a temperate outdoor environment on performance and physiology in the heat. Temp (Austin) 5(3):267–275. 10.1080/23328940.2018.1474672 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Waldron M, Fowler R, Heffernan S, Tallent J, Kilduff L, Jeffries O (2021) effects of heat acclimation and acclimatisation on maximal aerobic capacity compared to exercise alone in both thermoneutral and hot environments: a meta-analysis and meta-regression. Sports Med 51(7):1509–1525. 10.1007/s40279-021-01445-6 [DOI] [PMC free article] [PubMed] [Google Scholar]
- Wallace PJ, Hartley GL, Nowlan JG, Ljubanovich J, Sieh N, Taber MJ, Gagnon DD, Cheung SS (2024) Endurance capacity impairment in cold air ranging from skin cooling to mild hypothermia. J Appl Physiol 136(1):58–69. 10.1152/japplphysiol.00663.2023 [DOI] [PubMed] [Google Scholar]
- Warburton DE, Gledhill N, Quinney HA (2000) Blood volume, aerobic power, and endurance performance: potential ergogenic effect of volume loading. Clin J Sport Med 10(1):59–66. 10.1097/00042752-200001000-00011 [DOI] [PubMed] [Google Scholar]
- Wickham KA, Wallace PJ, Cheung SS (2021) Sex differences in the physiological adaptations to heat acclimation: a state-of-the-art review. Eur J Appl Physiol 121(2):353–367. 10.1007/s00421-020-04550-y [DOI] [PubMed] [Google Scholar]
- Willmott AGB, Gibson OR, James CA, Hayes M, Maxwell NS (2018) Physiological and perceptual responses to exercising in restrictive heat loss attire with use of an upper-body sauna suit in temperate and hot conditions. Temperature (Austin) 5(2):162–174. 10.1080/23328940.2018.1426949 [DOI] [PMC free article] [PubMed] [Google Scholar]
