Oxygen uptake kinetics from ramp work tests: variability of single test values

The variability in the estimation of the mean response time (MRT) of O2 uptake (VO2) kinetics from single ramp work rate exercise tests was examined in six repetitions by five fit subjects. Work rate increased at 50 W/min from a base line of 25 W to a work rate of 120% ventilatory threshold. Breath-by-breath data were analyzed by linear regression from 2 min after the onset of the ramp to the 120% work rate. Individual subjects showed approximately twofold differences in estimates of MRT; the coefficient of variation from individuals ranged from 18.5 to 29.3%. The MRT obtained as the mean from the individual repetitions did not differ from the MRT obtained from pooled within-subject data. Analysis of variance on the individual MRT estimates showed 53.9% of the variability was attributable to the slope of the regression, whereas only 2.4% could be attributed to baseline VO2. It was concluded that several repetitions of the ramp work rate tests should be pooled prior to estimation of kinetics parameters.     

Water immersion delays the oxygen uptake response to sitting arm-cranking in humans

We investigated the effect of central hypervolaemia during water immersion up to the xiphoid process on the oxygen uptake (VO2) and heart rate (HR) response to arm cranking. Seven men performed a 6-min arm-cranking exercise at an intensity requiring a VO2 at 80% ventilatory threshold both in air [C trial, 29 (SD 9) W] and immersed in water [WI trial, 29 (SD 11) W] after 6 min of sitting. The VO2 (phase 2) and HR responses to exercise were obtained from a mono-exponential fit [f(t) = baseline + gain x (1 - e(-(t-TD)/tau))]. The response was evaluated by the mean response time [MRT; sum of time constant (tau) and time delay (TD)]. No significant difference in VO2 and HR gains between the C and WI trials was observed [VO2 0.78 (SD 0.1) vs 0.80 (SD 0.2) l x min(-1), HR 36 (SD 7) vs 37 (SD 8) beats x min(-1), respectively]. Although the HR MRT was not significantly different between the C and WI trials [17 (SD 3), 19 (SD 8) s, respectively), VO2 MRT was greater in the WI trial than in the C trial [40 (SD 6), 45 (SD 6) s, respectively; P < 0.05]. Assuming no difference in VO2 in active muscle between the two trials, these results would indicate that an increased oxygen store and/or an altered response in muscle blood distribution delayed the VO2 response to exercise.     

Time domain analysis of oxygen uptake during pseudorandom binary sequence exercise tests.

Pseudorandom binary sequence (PRBS) exercise tests involve repeated switching between two work rates (WR) according to a computer-generated pattern. This paper presents an approach to analysis of O2 uptake (VO2) in the time domain. First, the autocorrelation function (ACF) of the input WR was recognized to be a triangular-shaped pulse that can be taken to be equivalent to a ramp increase followed by a ramp decrease in WR. Then the cross-correlation function of the input (WR) and the output (VO2) was treated as if it were the response to a triangular-shaped pulse. The cross-correlation function was analyzed by fitting a linear summation of the ramp form of a two-component exponential function to this triangular pulse. VO2 responses of eight subjects were obtained from two different PRBS tests, as well as step changes in WR. The first PRBS test consisted of 15 units, each 30 s in duration. Its ACF had a base width of 60 s. The ramp increase-ramp decrease model fit the data throughout the range of response. The second PRBS test had 63 units, each 5 s in duration; thus its ACF base width was 10 s. Again, the ramp model fit adequately. The data from the second PRBS test could be fit by the impulse form of the two-component exponential equation, although the fit in the first 30 s tended to be poorer. The time constants of VO2 dynamics estimated from step and PRBS tests were not significantly different. PRBS tests can be analyzed in the time domain, and the indicators of system dynamics reflect physiological properties similar to those investigated during step changes in WR.     

Estimate of mean tissue O2 consumption at onset of exercise in males

A mathematical model has been developed that permitted the calculation of the flow-weighted mean tissue O2 consumption (VO2T) at the onset of a step increase in work rate. From breath-by-breath measurements of alveolar O2 consumption (VO2A) and cardiac output (Q) by impedance cardiography and assumptions about the site of depletion of O2 stores, the rate of change in O2 stores (VO2s) was determined. The sum of VO2A + VO2s = VO2T. Six very fit males performed six repetitions of each of two step increases in work rate. STlo was a transition from rest to 100-W cycling; SThi was a transition from 100- to 200-W cycling. For each work rate transition, the responses of VO2A and Q were averaged over the six repetitions of each subject and the model was solved to yield VO2T. The responses of VO2A, VO2T, and Q after the increase in work rate were fit with a monoexponential function. This function included a time constant and time delay, the sum of which gave the mean response time (MRT). In the STlo test, the MRT of VO2A (24.9 +/- 1.1 s, mean +/- SE) was longer than that of VO2T (15.3 +/- 1.3 s) and of Q (16.5 +/- 6.5 s) (P less than 0.05). The MRT of VO2T and Q did not differ significantly. Also for SThi, the MRT of VO2A (34.4 +/- 3.3 s) was significantly longer than that of VO2T (30.0 +/- 3.4 s) (P less than 0.05). The MRT of VO2T and Q (30.3 +/- 5.5 s) were not significantly different at this work rate either.(ABSTRACT TRUNCATED AT 250 WORDS)     

VCO2 and VE kinetics during moderate- and heavy-intensity exercise after acetazolamide administration.

The effect of carbonic anhydrase inhibition with acetazolamide (Acz) on CO2 output (VCO2) and ventilation (VE) kinetics was examined during moderate- and heavy-intensity exercise. Seven men [24 +/- 1 (SE) yr] performed cycling exercise during control (Con) and Acz (10 mg/kg body wt iv) sessions. Each subject performed step transitions (6 min) in work rate from 0 to 100 W [below ventilatory threshold (VET)]. VE and gas exchange were measured breath by breath. The time constant (tau) was determined for exercise VET by using a three-component model (fit from the start of exercise). VCO2 kinetics were slower in Acz (VET, MRT = 75 +/- 10 s) than Con (VET, MRT = 54 +/- 7 s). During VET kinetics were faster in Acz (MRT = 85 +/- 17 s) than Con (MRT = 106 +/- 16 s). Carbonic anhydrase inhibition slowed VCO2 kinetics during both moderate- and heavy-intensity exercise, demonstrating impaired CO2 elimination in the nonsteady state of exercise. The slowed VE kinetics in Acz during exercise 相似文献   

Gas exchange responses to ramp exercise

In the present study, the system of oxygen uptake (VO2) during ramp function exercise protocol can be studied to provide information about the physiological mechanisms underlying the process. The values of maximal oxygen uptake (VO2max) and gas exchange threshold (GET) were reproducibly obtained using ramp test protocol. On the other hand, the determination of VO2/work rate should be restrict to below the exercise intensity of the GET. Therefore, ramp exercise test might be usage for determination of VO2max, GET and/or VO2/work rate (i.e., work efficiency). The data obtained in this study concerning the mean response time (MRT) suggests that the ramp test is not a linear, first-order system. Therefore, the ramp exercise test protocol is recommended for the determination of VO2max, GET and work efficiency, but not for MRT.     

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.     

Parameters of ventilatory and gas exchange dynamics during exercise

To determine the precise nonsteady-state characteristics of ventilation (VE), O2 uptake (VO2), and CO2 output (VCO2) during moderate-intensity exercise, six subjects each underwent eight repetitions of 100-W constant-load cycling. The tests were preceded either by rest or unloaded cycling ("0" W). An early component of VE, VO2, and VCO2 responses, which was obscured on any single test by the breath-to-breath fluctuations, became apparent when the several repetitions were averaged. These early responses were abrupt when the work was instituted from rest but were much slower and smaller from the 0-W base line and corresponded to the phase of cardiodynamic gas exchange. Some 20 s after the onset of the work a further monoexponential increase to steady state occurred in all three variables, the time constants of which did not differ between the two types of test. Consequently, the exponential behavior of VE, VO2, and VCO2 in response to moderate exercise is best described by a model that incorporates only the second phase of the response.     

Ramp work tests with three different beta-blockers in normal human subjects

The effects of beta-blockade on the responses of oxygen uptake (VO2), heart rate (HR) and blood lactate (La-) were examined during ramp cycle ergometer tests (50 W.min-1 ramp slope) in 8 healthy male volunteers. Each subject took placebo, or one of four different doses of three different beta-blockers (propranolol, metoprolol or oxprenolol) 2 h prior to each test for a total of 15 exercise tests. VO2 was measured breath-by-breath, HR was sampled once per breath, and La- was obtained every minute. Linear regression analysis was applied to VO2 and HR data to obtain the kinetic parameter total lag time (TLT) and a slope value. La- was analyzed by a continuous exponential model with the lactate slope index (LSI) being derived from the individual response curves. Submaximal exercise HR was significantly depressed at the baseline as well as during the ramp tests by beta-blockade. TLT for HR was significantly affected by beta-blockade, with a dose dependent shift from a placebo value of 16 to 26 s with placebo to a value of -40 to -60 s at the highest dose. Slope of HR was significantly depressed relative to placebo. VO2 kinetics assessed by TLT were not significantly affected by beta-blockade. This slope of the VO2 vs work rate relationship was significantly less than placebo only at the highest dose of beta-blocker. The LSI was not significantly affected by beta-blockade. In contrast with the clear impairment of HR response to exercise during beta-blockade, both the VO2 and La- responses appear to be relatively unaffected by beta-blockade during ramp exercise tests.(ABSTRACT TRUNCATED AT 250 WORDS)     

Cardiovascular and respiratory responses to slow ramp carotid sinus pressures in the dog

The respiratory and mean arterial pressure (MAP) responses to slow ramp pressure stimulation of carotid baroreceptors were compared in pentobarbital-anesthetized vagotomized dogs breathing 100% O2. Carotid sinus pressure (CSP) was raised from 50 (control) to 220 mmHg and then returned to control as linear ramps (+/- 1 mmHg/s) in isolated sinuses. MAP, heart rate (HR), ventilation (VE), frequency (f), and tidal volume (VT) were expressed as percent of control. The maximum difference between responses to positive and negative ramps at a given CSP (MAX) and the average difference (AVG) served as indicators of the hysteresis for each response. In 27 dogs MAP changed monotonically with varying CSP with insignificant (P = 0.27, MAX) or barely significant (P = 0.03, AVG) hysteresis, monotonic function being one that is continuously nondecreasing or continuously nonincreasing. Similar responses were obtained for HR. VE decreased as CSP increased, but the change was not monotonic. During negative ramp, VE increased back to control with an overshoot. Hysteresis for VE was pronounced (P less than 0.0001, both measures). The VE response was primarily determined by f; VT increased with CSP. To eliminate secondary respiratory effects due to alterations in MAP, in seven dogs similar experiments were performed after ganglionic blockade with hexamethonium. Hysteresis in VE and f persisted. To assess the role of changing arterial PCO2 (PaCO2) on VE, the CSP was held constant (after a ramp rise) at 140, 150, or 180 mmHg before reducing it at -1 mmHg/s to 50 mmHg; however, a significant hysteresis in VE was still observed. Further experiments, to eliminate secondary reflexes due to altered PaCO2, were performed in seven dogs after ganglionic blockade and paralysis with Flaxedil, with phrenic nerve activity as an indicator of ("neural") respiration. The hysteresis in VE and f were no longer significant. In summary, the results indicate that 1) slow ramp carotid baroreceptor stimulation elicits both VE and cardiovascular responses, the VE response showing a dramatically higher hysteresis than the cardiovascular responses; 2) the ventilatory hysteresis is partially explained by the secondary changes in PaCO2 and perhaps by cardiovascular variables; and 3) the central processing of the baroventilatory reflex appears to be rate sensitive at a slower rate of pressure change than that which causes rate sensitivity in the baropressure reflex.     

Delayed kinetics of respiratory gas exchange in the transition from prior exercise

The kinetics of oxygen uptake (VO2), carbon dioxide output (VCO2), and expired ventilation (VE) in the transition from rest or from prior exercise were studied in response to step increases in power output (PO). The data were modeled with a single-component exponential function incorporating a time delay (TD). Each subject exercised on four occasions. Test 1 was an incremental test for determination of ventilatory anaerobic threshold (AT). Step increase tests were rest to 80% of PO at AT (test 2), rest-40% AT (3a), 40-80% AT (3b), rest-40% AT (4a), and 40-120% AT (4b). Respiratory gas exchange was monitored by open-circuit techniques. The VO2 kinetics showed the time constant (tau) to be longer in the transitions from prior exercise [tests 3b and 4b were 60.6 +/- 10.8 (SD) and 79.2 +/- 17.4 s] than from rest (tests 2, 3a, and 4a were 37.8 +/- 7.2, 30.0 +/- 7.8, and 39.6 +/- 17.4 s). The mean response time (MRT = tau + TD) was also longer for these tests. Kinetic analysis for VCO2 showed a tendency for tau to be shorter for the tests from prior exercise, but neither tau nor tau + TD were significantly different between tests. In contrast to VCO2, VE kinetics showed a significantly longer tau + TD for test 3b (P less than 0.05) and test 4b (P less than 0.01). This study has shown the VO2 kinetics to be delayed when a given increment in PO occurred from prior exercise, whether the final PO was below or above the AT. Further, the dissociation of VCO2 and VE kinetics does not support a direct link between these two variables as the sole control factor in exercise hyperpnea.     

A comparison of the effect of external loading upon power output in stair climbing and running up a ramp

Previous studies have shown that external loading increases the power output measured during stair climbing. However, it was noted in an earlier study that stairtreads form mechanical contraints which limit the extent to which a subject can be externally loaded, and, thereby, make it impossible to observe maximal power output for this type of activity. The purpose of this study was to compare the effects of external loading upon power output when running up stairs or a ramp. Since a ramp is free of the mechanical constraints of stairtreads, it was felt that higher power output values would be achieved using the ramp, and that it would be possible to observe an asymptote in power output which could not be obtained for stair climbing. Seven male subjects performed maximal ramp and stair climbing tests under five experimental loading conditions (no external load, 10.1, 19.2, 24.2, and 29.2 kg). For the ramp, it was possible to employ a sixth loading condition of 34.2 kg. For stair climbing, the mean (+/- SD) power output values under the five experimental conditions were 16.6 +/- 0.7, 17.3 +/- 1.3, 18.5 +/- 1.0, 18.6 +/- 1.5, and 18.9 +/- 1.7 W X kg-1, respectively. In contrast, the mean (+/- SD) power output values observed while running up the ramp were 18.8 +/- 1.4, 19.9 +/- 1.6, 20.5 +/- 1.6, 20.1 +/- 2.1, 20.3 +/- 2.1, and 19.8 +/- 1.9 W X kg-1, respectively. At each experimental condition, the differences between the ramp and stairs was significant (P less than 0.05).(ABSTRACT TRUNCATED AT 250 WORDS)     

Kinetics of ventilation and gas exchange during supine and upright cycle exercise.

The dynamics of ventilation (VE), oxygen uptake (VO2), carbon dioxide output (VCO2), and heart rate (fc) were studied in 12 healthy young men during upright and supine exercise. Responses to maximal and to two different types of submaximal exercise tests were contrasted. During incremental exercise to exhaustion, the maximal work rate, VO2max, VEmax, fc,max, and ventilatory threshold were all significantly reduced in supine compared to upright exercise (P less than 0.01-0.001). Following step increases or decreases in work rate between 25 W and 105 W, both VO2 and VCO2 responded more slowly in supine than upright exercise. Dynamics were also studied in two different pseudorandom binary-sequence (PRBS) exercise tests, with the work rate varying between 25 W and 105 W with either 5-s or 30-s durations of each PRBS unit. In both of these tests, there were no differences caused by body position in the amplitude or phase shifts obtained from Fourier analysis for any observed variable. These data show that the body position alters the dynamic response to the more traditional step increase in work rate, but not during PRBS exercise. It is speculated that the elevation of cardiac output observed with supine exercise in combination with the continuously varying work-rate pattern of the PRBS exercise allowed adequate, perhaps near steady-state, perfusion of the working muscles in these tests, whereas at the onset of a step increase in work rate, greater demands were placed on the mechanisms of blood flow redistribution.     

Microwave-Assisted Extraction of Wax from Oily Sludge: An Experimental Study and its Process Variables Optimization Using Response Surface Methodology

The wax present in petroleum sludge, generated by refineries and at crude production sites, consists of paraffin hydrocarbons (C18–C36) known as paraffin wax and naphthenic hydrocarbons (C30–C60). The present study is aimed at the recovery of wax from petroleum oily sludge by microwave-assisted solvent extraction using a Toluene/MEK mixture and subsequently de-crystallizing the wax. The process variables affecting the microwave-assisted solvent extraction are optimized for recovery of wax. The simultaneous effects of process variables such as irradiation time (2–10 minutes), solvent to sludge ratio (40–80 wt%), reactant volume (100–300 ml), and microwave power (80–400 W) on the recovery of wax were evaluated. A central composite design and response surface methodology were used for the optimization of the extraction process. Based on the central composite design, quadratic models were developed to correlate the extraction process variables with the responses and the models were analyzed using appropriate statistical methods for analysis of variance. Optimization of process variables shows the maximum recovery of wax was about 79.57% at 300 ml of reactant volume with microwave power output of 400 W at 7.6 minutes of retention time with 56.56% of Toluene/MEK to sludge ratio.     

Similar level of impairment in exercise performance and oxygen uptake kinetics in middle-aged men and women with type 2 diabetes

The present study tested the hypothesis that the magnitude of the type 2 diabetes-induced impairments in peak oxygen uptake (Vo(2)) and Vo(2) kinetics would be greater in females than males in middle-aged participants. Thirty-two individuals with type 2 diabetes (16 male, 16 female), and 32 age- and body mass index (BMI)-matched healthy individuals (16 male, 16 female) were recruited. Initially, the ventilatory threshold (VT) and peak Vo(2) were determined. On a separate day, subjects completed four 6-min bouts of constant-load cycling at 80% VT for the determination of Vo(2) kinetics using standard procedures. Cardiac output (CO) (inert gas rebreathing) was recorded at rest, 30, and 240 s during two additional bouts. Peak Vo(2) (ml·kg(-1)·min(-1)) was significantly reduced in men and women with type 2 diabetes compared with their respective nondiabetic counterparts (men, 27.8 ± 4.4 vs. 31.1 ± 6.2 ml·kg(-1)·min(-1); women, 19.4 ± 4.1 vs. 21.4 ± 2.9 ml·kg(-1)·min(-1)). The time constant (s) of phase 2 (τ(2)) and mean response time (s) of the Vo(2) response (MRT) were slowed in women with type 2 diabetes compared with healthy women (τ(2), 43.3 ± 9.8 vs. 33.6 ± 10.0 s; MRT, 51.7 ± 9.4 vs. 43.5 ± 11.4s) and in men with type 2 diabetes compared with nondiabetic men (τ(2), 43.8 ± 12.0 vs. 35.3 ± 9.5 s; MRT, 57.6 ± 8.3 vs. 47.3 ± 9.3 s). The magnitude of these impairments was not different between males and females. The steady-state CO responses or the dynamic responses of CO were not affected by type 2 diabetes among men or women. The results suggest that the type 2 diabetes-induced impairments in peak Vo(2) and Vo(2) kinetics are not affected by sex in middle aged participants.     

The influence of natural diet composition, food intake level, and body size on ingesta passage in primates

An important component of digestive physiology involves ingesta mean retention time (MRT), which describes the time available for digestion. At least three different variables have been proposed to influence MRT in herbivorous mammals: body mass, diet type, and food intake (dry matter intake, DMI). To investigate which of these parameters influences MRT in primates, we collated data for 19 species from trials where both MRT and DMI were measured in captivity, and acquired data on the composition of the natural diet from the literature. We ran comparative tests using both raw species values and phylogenetically independent contrasts. MRT was not significantly associated with body mass, but there was a significant correlation between MRT and relative DMI (rDMI, g/kg(0.75)/d). MRT was also significantly correlated with diet type indices. Thus, both rDMI and diet type were better predictors of MRT than body mass. The rDMI-MRT relationship suggests that primate digestive differentiation occurs along a continuum between an "efficiency" (low intake, long MRT, high fiber digestibility) and an "intake" (high intake, short MRT, low fiber digestibility) strategy. Whereas simple-stomached (hindgut fermenting) species can be found along the whole continuum, foregut fermenters appear limited to the "efficiency" approach.     

Deciphering the metabolic and mechanical contributions to the exercise-induced circulatory response: insights from eccentric cycling

Metabolic demand and muscle mechanical tension are closely coupled during exercise, making their respective drives to the circulatory response difficult to establish. This coupling being altered in eccentric cycling, we implemented an experimental design featuring eccentric vs. concentric constant-load cycling bouts to gain insights into the control of the exercise-induced circulatory response in humans. Heart rate (HR), stroke volume (SV), cardiac output (Q), oxygen uptake (V(.-)(O(2))), and electromyographic (EMG) activity of quadriceps muscles were measured in 11 subjects during heavy concentric (heavy CON: 270 +/- 13 W; V(.-)(O(2)) = 3.59 +/- 0.20 l/min), heavy eccentric (heavy ECC: 270 +/- 13 W, V(.-)(O(2)) = 1.17 +/- 0.15 l/min), and light concentric (light CON: 70 +/- 9 W, V(.-)(O(2)) = 1.14 +/- 0.12 l/min) cycle bouts. Using a reductionist approach, the circulatory responses observed between heavy CON vs. light CON (difference in V(.-)(O(2)) and power output) was ascribed either to metabolic demand, as estimated from heavy CON vs. heavy ECC (similar power output, different V(.-)(O(2))), or to muscle mechanical tension, as estimated from heavy ECC vs. light CON (similar V(.-)(O(2)), different power output). 74% of the Q response was determined by the metabolic demand, also accounting for 65% and 84% of HR and SV responses, respectively. Consequently, muscle mechanical tension determined 26%, 35%, and 16% of the Q, HR, and SV responses, respectively. Q was significantly related to V(.-)(O(2)) (r(2) = 0.83) and EMG activity (r(2) = 0.82; both P < 0.001). These results suggest that the exercise-induced circulatory response is mainly under metabolic control and support the idea that the level of muscle activation plays a role in the cardiovascular regulation during cycle exercise in humans.     

Phase I dynamics of cardiac output, systemic O2 delivery, and lung O2 uptake at exercise onset in men in acute normobaric hypoxia

We tested the hypothesis that vagal withdrawal plays a role in the rapid (phase I) cardiopulmonary response to exercise. To this aim, in five men (24.6+/-3.4 yr, 82.1+/-13.7 kg, maximal aerobic power 330+/-67 W), we determined beat-by-beat cardiac output (Q), oxygen delivery (QaO2), and breath-by-breath lung oxygen uptake (VO2) at light exercise (50 and 100 W) in normoxia and acute hypoxia (fraction of inspired O2=0.11), because the latter reduces resting vagal activity. We computed Q from stroke volume (Qst, by model flow) and heart rate (fH, electrocardiography), and QaO2 from Q and arterial O2 concentration. Double exponentials were fitted to the data. In hypoxia compared with normoxia, steady-state fH and Q were higher, and Qst and VO2 were unchanged. QaO2 was unchanged at rest and lower at exercise. During transients, amplitude of phase I (A1) for VO2 was unchanged. For fH, Q and QaO2, A1 was lower. Phase I time constant (tau1) for QaO2 and VO2 was unchanged. The same was the case for Q at 100 W and for fH at 50 W. Qst kinetics were unaffected. In conclusion, the results do not fully support the hypothesis that vagal withdrawal determines phase I, because it was not completely suppressed. Although we can attribute the decrease in A1 of fH to a diminished degree of vagal withdrawal in hypoxia, this is not so for Qst. Thus the dual origin of the phase I of Q and QaO2, neural (vagal) and mechanical (venous return increase by muscle pump action), would rather be confirmed.     

The relationship between absolute disparity and ocular vergence

The relationship between disparity and ocular vergence was investigated under closed-loop as well as under open-loop viewing conditions. First we examined whether vergence responded similarly to disparity presented under open-loop and closed-loop conditions. Similar response were observed in both conditions. The direct relationship between disparity and vergence was examined by presenting constant disparities between 0.2° and 4° under open-loop viewing conditions. Such vergence responses are described as the outputs of first-order low-pass filters with different filter characteristics for each amplitude of disparity. By analyzing the latency of vergence responses induced by constant disparities with help of the transfer function of disparitycontrolled vergence, the time delay of disparity processing in the vergence loop was estimated. We suggested that the time delay was approximately between 80 and 120 ms instead of 160 ms as is generally assumed. The relationship between the rate of disparity change and vergence was examined by comparing responses to ramp and stepwise changes in target vergence. From the similar responses to ramp and staircase changes in disparity we concluded that vergence is not sensitive to the velocity of target vergence as such. On the basis of these findings we developed a model of disparity-controlled vergence. In this model disparity is processed through several parallel, imperfect integrators with slightly different low-pass filter characteristics, each of them susceptible to a limited range of disparities. Gains as well as phase lags of vergence responses to sinusoidal disparities are accurately simulated by this model. As a consequence of the limited working range of the low-pass filters, the model correctly simulates the alterations of fast and slow phases in response to step and ramps of target vergence, which are characteristic of real vergence responses.     

The dynamic nonlinear behavior of fly photoreceptors evoked by a wide range of light intensities.

Fly photoreceptor cells were stimulated with steps of light over a wide intensity range. First- and second-order Volterra kernels were then computed from sequences of combined step responses. Diagonal values of the second-order Volterra kernels were much greater than the off-diagonal values, and the diagonal values were roughly proportional to the corresponding first-order kernels, suggesting that the response could be approximated by a static nonlinearity followed by a dynamic linear component (Hammerstein model). The amplitudes of the second-order kernels were much smaller in light-adapted than in dark-adapted photoreceptors. Hammerstein models constructed from the step input/output measurements gave reasonable approximations to the actual photoreceptor responses, with light-adapted responses being relatively better fitted. However, Hammerstein models could not account for several features of the photoreceptor behavior, including the dependence of the step response shape on step amplitude. A model containing an additional static nonlinearity after the dynamic linear component gave significantly better fits to the data. These results indicate that blowfly photoreceptors have a strong early gain control nonlinearity acting before the processes that create the characteristic time course of the response, in addition to the nonlinearities caused by membrane conductances.