Diurnal variations in ventilatory and cardiorespiratory responses to submaximal treadmill exercise in females

Diurnal variations in ventilatory and cardiorespiratory responses to submaximal treadmill exercise were analysed in 11 eumenorrhoeic women and in 10 women using monophasic oral contraceptives. Subjects performed submaximal treadmill exercise at three intensities averaging 7, 8, and 9 km x h(-1), each for 4 min at 0800, 1300 and 1700 hours, assigned randomly on 3 separate days. Rectal temperature was measured before (T(rec(b))) and after (T(rec(a))) exercise. Cardiac frequency (f(c)), ventilation (V(E)), oxygen uptake (VO(2)), carbon dioxide output (VCO(2)), and respiratory exchange ratio (R) were assessed in the last minute of each stage of the exercise. Both T(rec(b)) and T(rec(a)) increased from 0800 to 1700 hours (P < 0.001). For a given submaximal work rate, VO(2) and VCO(2) were higher in the afternoon compared to the morning. Similarly, R was increased at 1700 hours compared to 0800 hours during the recovery period following exercise (P < 0.05). However, V(E) did not vary significantly during the day at any of the running intensities. No significant interactions (group x time of day) were observed in any of the studied parameters. In contrast to ventilation, the VO(2) and VCO(2) of the females during submaximal exercise were both affected by the time of day, without any differences between eumenorrhoeic women and users of oral contraceptives.     

Inflammatory cytokines and metabolic responses to high-intensity intermittent training: effect of the exercise intensity

To examine the effects of two high-intensity intermittent training (HIIT) programs of varying intensities (100% vs. 110% of maximal aerobic velocity [MAV]) on metabolic, hormonal and inflammatory markers in young men. Thirty-seven active male volunteers were randomly assigned into: HIIT experimental groups (100% MAV [EG₁₀₀, n = 9] and 110% MAV [EG₁₁₀, n = 9]) and a control groups (CG₁₀₀, n = 9 and CG₁₁₀, n = 9). Particpants performed high intesity intermittent exercise test (HIIE) at 100% or 110% MAV. Venous blood samples were obtained before, at the end of HIIE and at 15 min of recovery, and before and after 8 weeks of HIIT programs. After training, Glucose was lower (p < 0.01) in EG₁₀₀ (d = 0.72) and EG₁₁₀ (d = 1.20) at the end of HIIE, and at 15 min recovery only in EG₁₁₀ (d = 0.95). After training, Insulin and Cortisol were lower than before training in EG₁₀₀ and EG₁₁₀ at the end of HIIE (p < 0.001). After HIIT, IL-6 deceased (p < 0.001) in EG₁₀₀ (d = 1.43) and EG₁₁₀ (d = 1.56) at rest, at the end of HIIE (d = 1.03; d = 1.75, respectively) and at 15 min of recovery (d = 0.88;d = 1.7, respectively). This decrease was more robust (p < 0.05) in EG₁₁₀ compared to EG₁₀₀. After HIIT, TNF-α deceased (p < 0.001) in EG₁₀₀ (d = 1.43) and EG₁₁₀ (d = 0.60) at rest, at the end of HIIE (0.71 < d < 0.98) and at 15 min of recovery (0.70 < d < 2.78). HIIT with 110% MAV is more effective in young males on the improvements of some metabolic (Glucose), hormonal (Cortisol) and inflammatory (IL-6) markers at rest, at the end of HIIE and 15 min of recovery than training at 100 % MAV.     

A comparison of skeletal muscle oxygenation and fuel use in sustained continuous and intermittent exercise

In this study we compared substrate oxidation and muscle oxygen availability during sustained intermittent intense and continuous submaximal exercise with similar overall (i.e. work and recovery) oxygen consumption (VO2). Physically active subjects (n = 7) completed 90 min of an intermittent intense (12 s work:18 s recovery) and a continuous submaximal treadmill running protocol on separate days. In another experiment (n = 5) we compared oxygen availability in the vastus lateralis muscle between these two exercise protocols using near-infrared spectroscopy. Initially, overall VO(2) (i.e. work and recovery) was matched, and from 37.5 min to 67.5 min of exercise was similar, although slightly higher during continuous exercise (8%; P < 0.05). Energy expenditure was constant (22.5-90 min of exercise) and was not different in intermittent intense [0.81 (0.01) kJ x min(-1). kg(-1)] and continuous submaximal [0.85 (0.01) kJ x min(-1) x kg(-1)] exercise. Overall exercise intensity, represented as a proportion of peak aerobic power (VO2(peak)), was 68.1 (2.5)% VO2(peak) and 71.8 (1.8)% VO2(peak) for intermittent and continuous exercise protocols, respectively. Fat oxidation was almost 3 times lower (P < 0.05) and carbohydrate oxidation was approximately 1.2 times higher (P < 0.05) during intermittent compared to continuous exercise, despite the same overall energy expenditure. Capillary plasma lactate was constant from 15 to 90 min of exercise, and pyruvate was constant from 15 to 75 min, although both were higher (P < 0.0001, lactate; P < 0.001, pyruvate) during intermittent [5.05 (0.28) mM, 200 (7) microM, respectively] compared to continuous exercise [2.41 (0.10) mM, 114 (4) microM, respectively]. There was no difference between protocols for either plasma glycerol or non-esterified fatty acids. The decrease in muscle oxygenation during work periods of intermittent exercise resulted in a lower nadir oxygenation [54.62 (0.41)%] compared to continuous exercise [58.82 (0.21)%, P < 0.001]. The decline in oxygenation was correlated with treadmill speed (r = 0.72; P < 0.05). These results show a difference in substrate utilisation and muscle oxygen availability during sustained intermittent intense and continuous submaximal exercise, despite a similar overall VO(2) and identical energy expenditure.     

Growth hormone responses to growth hormone-releasing hormone (1-29)-NH2 and a D-Ala2 analog in normal men

Synthetic analogs of growth hormone-releasing hormone, GHRH(1-29)-NH2 and D-Ala2 GHRH(1-29)-NH2 were administered as a bolus intravenous injection to five normal men in a dose range of 0.015 to 0.5 micrograms/kg body weight. Vehicle only was administered in a control study. Peak responses to GHRH analogs occurred at 15 or 30 min. An increase in the integrated plasma growth hormone (GH) response was observed at each dose. The dose-response curve of GHRH(1-29)-NH2 indicated that it has a similar molar potency to GHRH(1-40) and GHRH(1-44). The potency of D-Ala2 GHRH(1-29)-NH2 was approximately twice that of GHRH(1-29)-NH2. Neither analog affected blood levels of PRL, TSH, LH, FSH, ACTH, insulin, glucagon, glucose, cortisol, free thyroxine, and free triiodothyronine. No side effects were noted other than transient flushing with the highest dose administered. The findings demonstrate GHRH(1-29)-NH2 and its D-Ala2 analog are potent stimulators of GH release and have potential application in clinical medicine.     

Plasma vasopressin, growth hormone and ACTH responses to static handgrip in healthy subjects

Ten healthy male subjects took part in the study. They performed three consecutive bouts of static handgrip at 30% of maximal voluntary contraction (MVC), using two hands alternately and without rest intervals. Blood pressure was measured every 30 s and ECG was recorded continuously. Blood samples for arginine vasopressin (AVP), growth hormone (GH), adrenocorticotrophic hormone (ACTH) and cortisol determinations were taken at rest, after each exercise bout, as well as at 10 and 30 min after the last one. During the whole period of exercise (9 min) blood pressure and heart rate were elevated. The effort caused a significant increase in the plasma AVP concentration. In the majority of subjects the peak values occurred after the first or second exercise bout and were followed by a rapid decline of the hormone concentration. Changes in GH, ACTH and cortisol concentrations were insignificant; however, in seven of the ten subjects, considerably elevated plasma GH levels were found at the end of the third exercise bout and/or 10 min after its cessation.     

Acute hormonal responses to heavy resistance exercise in younger and older men

The purpose of this investigation was to examine the acute responses of several hormones [total and free testosterone (TT and FT, respectively), adrenocorticotropic hormone (ACTH), cortisol (C), growth hormone (GH), and insulin (INS)] to a single bout of heavy resistance exercise (HRE). Eight younger [30-year (30y) group] and nine older [62-year (62y) group] men matched for general physical characteristics and activity levels performed four sets of ten repetitions maximum (RM) squats with 90 s rest between sets. Blood samples were obtained from each subject via an indwelling cannula with a saline lock pre-exercise, immediately post-exercise (IP), and 5, 15 and 30 min post-exercise. Levels of TT, FT, ACTH, C and lactate significantly increased after HRE for both groups. Pre-HRE pairwise differences between groups were noted only for FT, while post-HRE pairwise differences were found for TT, FT, GH, glucose and lactate. Area under the curve analysis showed that the 30y group had a significantly higher magnitude of increase over the entire recovery period (IP, 5, 15, and 30 min post-exercise) for TT, FT, ACTH and GH. Few changes occurred in the INS response with the only change being that the 62y group demonstrated a decrease IP. Lactate remained elevated at 30 min post-HRE. This investigation demonstrates that age-related differences occur in the endocrine response to HRE, and the most striking changes appear evident in the FT response to HRE in physically active young and older men. Accepted: 11 June 1997     

Seasonal changes in ovarian steroid hormone concentrations in the large hairy armadillo (Chaetophractus villosus) and the crying armadillo (Chaetophractus vellerosus)

Knowledge of armadillo reproductive physiology is essential for developing ex situ and in situ assisted reproductive techniques for propagating and/or controlling populations of these animals. The present study included assessment of fecal sex steroids by radioimmunoassay, determining reproductive status via monitoring ovarian activity (in the wild) and therefore reproductive status, in wild females of the large hairy armadillo (Chaetophractus villosus) and the crying armadillo (Chaetophractus vellerosus) in the southern hemisphere. Plasma and fresh fecal progesterone concentrations were not significantly correlated in either species. However, in both species, there was a significant positive correlation between plasma progesterone and dry fecal progesterone concentrations (r = 0.82, P < 0.05 and r = 0.60, P < 0.05, respectively). Dry fecal progesterone and estradiol concentrations were measured in one captive C. villosus (average baseline progesterone and estradiol concentrations 28.72 ± 11.75 ng/g dry feces and 3.04 ± 0.80 ng/g dry feces, respectively) and one captive C. vellerosus (average baseline progesterone and estradiol concentrations 14.05 ± 3.03 ng/g dry feces and 3.46 ± 1.20 ng/g dry feces, respectively) to detect hormonal peaks over 1 y; these occurred from late fall to early summer. Feces from wild C. villosus and C. vellerosus were also collected over 1 y to determine progesterone peaks, which occurred in winter and spring in both species (with no peaks during the summer or fall). Accordingly, C. villosus and C. vellerosus had a seasonal reproductive pattern. The significant correlations between dry fecal and plasma progesterone concentrations validated this method for monitoring reproductive status in these species.     

The influence of cold stress and a 36- h fast on the physiological responses to prolonged intermittent walking in man

In a previous study, rectal temperature (T _re) was found to be lower, and oxygen consumption (V˙O₂) and the respiratory exchange ratio (R) were higher in a cold (+5°C), wet and windy environment (COLD), compared with a thermoneutral environment during intermittent walking at ≈30% of peak V˙O₂ (Weller AS, Millard CE, Stroud MA et al. Am J Physiol 272:R226–R233, 1997). The aim of the present study was to establish whether these cold-induced responses are influenced by prior fasting, as impaired thermoregulation has been demonstrated in cold-exposed, resting men following a 48-h fast. To address this question, eight men attempted a 360-min intermittent (15 min rest, 45 min exercise) walking protocol under COLD conditions on two occasions. In one condition, the subjects started the exercise protocol ≈120 min after a standard meal (FED/COLD), whereas in the other the subjects had fasted for 36 h (FASTED/COLD). The first two exercise periods were conducted at a higher intensity (HIGHER, 6 km · h⁻¹ and 10% incline), than the four subsequent exercise periods (LOW, 5 km · h⁻¹ and 0% incline). There was no difference in the time endured in FED/COLD and FASTED/COLD. In FASTED/COLD com pared with FED/COLD, R was lower during HIGHER and LOW, and T _re was lower during LOW, whereas there was no difference in V˙O₂, mean skin temperature and heart rate. Therefore, although the 36-h fast impaired temperature regulation during intermittent low-intensity exercise in the cold, wet and windy environment, it was unlikely to have been the principal factor limiting exercise performance under these experimental conditions. Accepted: 26 August 1997     

Influence of training on the response to exercise of adrenocorticotropin and growth hormone plasma concentrations in human swimmers

The aim of the present study was to evaluate the response of adrenocorticotropin ([ACTH]) and growth hormone ([GH]) concentrations to a typical aerobic swimming set during a training season. Nine top-level male endurance swimmers (age range 17–23 years) were tested during three training sessions occurring 6, 12 and 18 weeks after the beginning of the season. During each session, after a standard warm-up, the swimmers performed a training set of 15 × 200-m freestyle, with 20 s of rest between repetitions, at a predetermined individual speed. Blood samples were collected before warm-up and at the end of the training set. A few days before each session, the individual swimming velocity corresponding to the 4 mmol · l⁻¹ blood lactate concentration (v ₄) was assessed as a standard of aerobic performance. Aerobic training affected v ₄ levels, which were highest 18 weeks after the beginning of the season; at the same time, while [ACTH] response was attenuated, [GH] response was enhanced. These results could be considered as adaptations to the exercise intensity. In our training programme, these adaptations seemed to have occurred between the 12th and 18th weeks of the training season. Accepted: 21 April 1998     

Growth hormone binding and stimulation of amino acid uptake in epithelial cells of rat ventral prostate

The binding of [125I]-human growth hormone (hGH) was studied in epithelial cells isolated from rat ventral prostate. Binding and degradation were dependent on time and temperature. The effect of a lysosomotropic agent suggested internalization and lysosomal degradation of the hormone. Dissociation and stoichiometric studies indicated the existence of a single class of GH receptors with a Kd of 0.7 nM and a binding capacity of 46 fmol hGH bound mg-1 cell protein. The receptor appeared to possess a somatotrophic nature since lactogenic hormones such as human placental lactogen and rat prolactin exhibited a very low degree of competition (whereas a variety of unrelated hormones and neuropeptides showed no effect). GH-stimulated leucine uptake by the cells in a time- and dose-dependent manner, half maximal effect being observed at 0.32 nM GH thus suggesting a direct relationship with the binding step.     

Comparisons of serum testosterone and corticosterone between exercise training during normoxia and hypobaric hypoxia in rats

The purpose of this study was to compare the effects of continuous and intermittent exercise training on serum testosterone [T] and corticosterone concentrations [Cort] during normoxia and hypobaric hypoxia. Male rats swam with loads of 3% (normoxia) or 2.25% (462 mmHg) body mass for 60 min in the continuous training groups, and 15 min separated by a 7-min rest  × 4, with 60-min total exercise duration in the intermittent training groups, 5␣days · week⁻¹ for 6 weeks. Serum [T] were measured at␣rest and following exercise after 6 weeks of training. Serum [Cort] were measured immediately after an acute period of exercise or after 6 weeks of training at rest and following exercise. Continuous exercise induced decreases in [T] under both conditions. Intermittent exercise showed a tendency to increase [T] during normoxia, but caused a suppression during hypobaric hypoxia. The [Cort] was elevated by a similar margin after an acute period of exercise during both conditions. After 6 weeks of training, however, [Cort] increased slightly after exercise during normoxia. A lower resting [Cort], which was increased after exercise, was found in the training groups during hypoxia. No relevant relationship was found between the behaviours of [T] and [Cort] after exercise during either conditions. Accepted: 20 April 1998     

Acute hormone responses to heavy resistance lower and upper extremity exercise in young versus old men

Acute hormone responses of growth hormone (GH), total and free testosterone (TT and FT) and cortisol (C) to heavy resistance isometric exercise were examined in ten young men [YM 26.5 (SD 4.8) years] and ten old men [OM 70.0 (SD 3.7) years]. Loading conditions of the same relative intensity were created for the lower and upper extremity actions separately as well as for both of them together – lower extremity exercise (LE; knee extension), upper extremity exercise (UE; bench press extension), and lower and upper extremity exercise (LUE) performed simultaneously in a seated position. Single voluntary maximal isometric actions lasting for 5 s were performed repeatedly for ten repetitions (with a recovery of 5 s) for a total of four sets. The recovery time between the sets was 1 min. Each exercise led to large acute decreases in maximal isometric force in both YM (P < 0.001) and OM (P < 0.001) ranging from 41% to 26% with no significant differences between the groups. Serum GH concentrations increased in both YM (P < 0.05–0.01) and OM (P < 0.05) but the postexercise value in YM during LE was greater (P < 0.05) than for OM. The TT increased (P < 0.01–0.001) in YM in all three exercises, while in OM the increase occurred only during LE (P < 0.01). The exercises led to increases in FT in YM (P < 0.05 for LE and LUE), while in OM the increase occurred only during LUE (P < 0.05). The pre and postexercise FT were greater in YM (P < 0.001) than in OM. No significant changes occurred in C either in YM or in OM. The blood lactate concentrations increased during the exercises in both YM (P < 0.001) and OM (P < 0.05–0.001) but the postexercise values during LE and LUE in YM were greater (P < 0.05) than in OM. The present data would indicate that the responses of GH, TT and FT to heavy resistance isometric exercise are lowered with increasing age. The reduced acute hormone response together with the lowered basal values in FT in the older men compared to the young men may indicate decreased anabolic effects on muscles and may explain in part the loss of muscle mass and strength associated with aging. Accepted: 18 August 1997     

Growth hormone kinetics during adaptation to a hyperosmotic environment in rainbow trout

Summary To clarify the role of growth hormone (GH) in salmonids during seawater (SW) adaptation, we examined GH kinetics in chronically cannulated rainbow trout, weighing about 1 kg. When trout were transferred from fresh water (FW) to 75% SW, plasma chloride concentration was normalized within 1 week. Plasma GH concentration increased significantly 2 days after transfer and decreased to the initial level thereafter. Metabolic clearance rate (MCR) and secretion rate (SR) of GH were calculated from the plasma levels of GH measured by radioimmunoassay after intra-arterial injection of recombinant chum salmon GH. 4 days after transfer to 75% SW, both MCR and SR increased to levels five times higher than those in FW, and returned to the FW levels after 3–4 weeks. In rainbow trout GH seems to be involved in the development of hypoosmoregulatory mechanisms, especially during the early phase of adaptation.     

缓释型促性腺激素释放激素与生长激素治疗儿童中枢性性早熟的疗效观察

目的用生长激素与促性腺激素释放激素类似物(Gonadotropin-releasing hormone analogues,GHA)联合治疗中枢性性早熟女性患儿对其最终成人身高的影响.方法生长激素(GH)与促性腺激素释放激素类似物(GHA)联合治疗4例中枢性性早熟女性患儿半年,对比治疗前后患儿的第二性征,骨龄发育,性激素及最终成人身高的变化.结果第二性征的发育停止,骨龄发育被控制,实际生活年龄与骨年龄的比值提高(平均0.79→0.84);血LH对促性腺激素释放激素的反应及血浆雌激素水平平均已降至青春期前,分别为(平均25.79±10.60mlu/ml→1.13±0.21mlu/ml)及(平均64.87±27.51pg/ml→3.03±1.87pg/ml);预测最终成人身高增加(平均149.60±4.31cm→156.75±3.84cm)差异具有显著性(P＜0.05).结论生长激素与GHA联合治疗中枢性性早熟患儿,不仅能抑制第二性征发育,而且能有效改善最终成人身高,无任何毒付作用.     

Influence of the menstrual cycle and oral contraceptives on thermoregulatory responses to exercise in young women

Thermoregulatory responses to exercise in relation to the phase of the menstrual cycle were studied in ten women taking oral contraceptives (P) and in ten women not taking oral contraceptives (NP). Each subject was tested for maximal aerobic capacity ( ) and for 50% exercise in the follicular (F) and luteal (L) phases of the menstrual cycle. Since the oral contraceptives would have prevented ovulation a quasi-follicular phase (q-F) and a quasi-luteal phase (q-L) of the menstrual cycle were assumed for P subjects. Exercise was performed on a cycle ergometer at an ambient temperature of 24° C and relative air humidity of 50%. Rectal (T _re), mean skin ( ), mean body ( ) temperatures and heart rate (f _c) were measured. Sweat rate was estimated by the continuous measurement of relative humidity of air in a ventilated capsule placed on the chest, converted to absolute pressure (PH₂O_chest). Gain for sweating was calculated as a ratio of increase inPH₂O_chest to the appropriate increase inT _re for the whole period of sweating (G) and for unsteady-state (G_u) separately. The did not differ either between the groups of subjects or between the phases of the menstrual cycle. In P, rectal temperature threshold for sweating (T _{re, td}) was 37.85° C in q-L and 37.60° C in q-F (P < 0.01) and corresponded to a significant difference fromT _re at rest. TheT _re, andf _c increased similarly during exercise in q-F and q-L. No menstrual phase-related differences were observed either in the dynamics of sweating or in G. In NP,T _{re, td} was shorter in L than in F (37.70 vs 37.47° C,P<0.02) with a significantly greater value fromT _re at rest. The dynamics and G for sweating were also greater in L than in F. The G_u was 36.8 versus 16.6 kPa · ° C^–1 (P<0.01) while G was 6.4 versus 3.8 kPa · ° C^–1 (P<0.05), respectively. TheT _re, andf _c increased significantly more in phase F than in phase L. It was concluded that in these women performing moderate exercise, there was a greater temperature threshold and larger gains for sweating in phase L than in phase F. Intake of oral contraceptives reduced the differences in the gains for sweating making the thermoregulatory responses to exercise more uniform.     

Different thermal sensitivity to exercise and cold in men and women

Rectal temperature gain for sweating and shivering was investigated in men (M) and women during exercise and cold exposure. Women, taking and not taking oral contraceptives, were tested during follicular (F) and luteal (L) phases of the menstrual cycle. The gains for sweating (M, 4.85; F, 5.00; L, 12.25 kPa °C⁻¹) and shivering (M, 6.50; F, 2.75; L, 2.00 ml kg min⁻¹ °C⁻¹) differed between sexes, as well as between F and L, and were affected by the oral contraceptives (5.45 and 8.00 kPa °C⁻¹, P<0.01, and 6.00 and 4.25 ml kg min⁻¹ °C⁻¹, P<0.05, respectively).     

Growth hormone induces eNOS expression and nitric oxide release in a cultured human endothelial cell line

Growth hormone deficiency is linked to cardiovascular disease and particularly increased peripheral vascular resistance. Surprisingly, its role in endothelial nitric oxide (NO) synthetase (eNOS) regulation and NO release is basically unknown. We therefore studied the effects of different doses of somatotropin in cultures of a human endothelial cell line (EAhy926). We investigated expression and activity of eNOS, as well as other target genes known to be deregulated in cardiovascular disease including E-selectin and the lectin-like oxidized low density lipoprotein receptor. Treatment of cultured human endothelial cells with somatotropin resulted in significant (P<0.05) increases of eNOS gene and protein expression, as well as NO release, whereas production of intracellular reactive oxygen species was significantly reduced, at the highest somatotropin dose level. The enhanced eNOS gene/protein expression and enzyme activity correlate well. Our findings are suggestive for a novel role of growth hormone in endothelial biology, and particularly NO production.     

Growth hormone and insulin-like growth factor I concentrations in bulls of various growth hormone genotypes

A leucine/valine substitution at amino acid position 127 was identified by the polymerase chain reaction and restriction fragment length polymorphism in the bovine growth hormone gene. Genotyping was performed in 84 AI bulls of three different breeds, in which plasma concentrations of growth hormone (GH) and insulin-like growth factor I (IGF-1) were also measured. Gene frequencies of variants L (leucine) und V (valine) were 0.80/0.20 (Black and White), 0.90/0.10 (Brown), 0.71/0.29 (Simmental). Hormone concentrations were measured during different physiological conditions (normal feeding, fasting, realimentation) in the majority of animals. Generally, genotype LL was associated with higher concentrations of GH than LV. This difference was significant in Black and White bulls (P < 0.05). In contrast, IGF-1 concentrations were higher in LV than in LL animals. This was most pronounced in mature, realimented Simmental bulls. We conclude that the various GH alleles influence the circulating concentrations of GH and IGF-1.     

Patterns of intermittent ventilation and responses to perfusing gas mixtures in quiescent Blaberus craniifer

ABSTRACT. Patterns of intermittent ventilation were recorded by means of long electromyogram wires from quiescent Blaberus craniifer (Burmeister) buried in vermiculite. While buried, cockroaches were subjected to perfusion with various mixtures of CO₂ in air and of oxygen in nitrogen. Quiescent cockroaches in air ventilated for mean periods of 138 s in cycles of 720s duration, but much variability occurred within and between cockroaches. Mild hypercapnia or hypoxia shortened the overall cycle time while more severe treatment caused the cycle to be replaced by continual pumping. Intermittent ventilation persisted in decapitated insects but the threshold of the response to hypoxia or hypercapnia was elevated. Prevailing gas tensions normally determine the frequency and duration of each phase.