Simplified models for gas exchange in the human lungs

This paper presents a hierarchy of models with increasing complexity for gas exchange in the human lungs. The models span from a single compartment, inflexible lung to a single compartment, flexible lung with pulmonary gas exchange. It is shown how the models are related to well-known models in the literature. A long-term purpose of this work is to study nonlinear phenomena seen in the cardio-respiratory system (for example, synchronization between ventilation rate and heart rate, and Cheyne-Stokes respiration). The models developed in this paper can be regarded as the controlled system (plant) and provide a mathematical framework to link between "molecular-level", and "systems-level" models. It is shown how changes in molecular level affect the alveolar partial pressure. Two assumptions that have previously been made are re-examined: (1) the hidden assumption that the air flow through the mouth is equal to the rate of volume change in the lungs, and, (2) the assumption that the process of oxygen binding to hemoglobin is near equilibrium. Conditions under which these assumptions are valid are studied. All the parameters in the models, except two, are physiologically realistic. Numerical results are consistent with published experimental observations.     

Nitrogen application improves gas exchange characteristics and chlorophyll fluorescence in maize hybrids under salinity conditions

The understanding of crop physiological responses to salinity stress is of paramount importance for selection of genotypes with improved tolerance to this stress. Maize (Zea mays L.) hybrids Pioneer 32B33 and Dekalb 979 were grown in pots and subjected to three levels of salinity under four nitrogen levels to determine the role of nitrogen under saline conditions. Salinity stress effects on gas exchange characteristics and chlorophyll fluorescence of maize hybrids were evaluated under semi-controlled conditions. Under salinity stress, the changes in the net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) were similarly directed: all decreased and were lower than in control at the higher salinity level (10 dS/m). Water use efficiency was increased with increasing salinity since transpiration was stronger depressed by salt than photosynthesis. Plants subjected to the lower level of salinity did not differ from control in tested characteristics. Nitrogen application ameliorated the effects of salinity.     

昆虫不连续气体交换

许多昆虫呼吸时气体交换是不连续的循环式进行的。根据气门开闭,一个典型的不连续气体交换循环（discontinuous gas exchangecycle, DGC）可以明显分为3个阶段: 关闭阶段,极少或没有气体交换;颤动阶段,气门迅速微开和关闭,O₂进入气管,少量CO₂释放;最后是开放阶段,大量的CO₂释放。该文综述了DGC特征及昆虫活动、温度、体重对DGC的影响,并讨论了DGC与呼吸失水、缺氧或高CO₂浓度环境有关的进化适应意义。     

Cutaneous gas exchange in snakes

Summary Cutaneous aquatic gas exchange and pulmonary gas exchange have been compared in an aquatic snakeAchrochordus javanicus and the terrestrial snakeConstrictor constrictor.Gas exchange was measured by closed respirometry with the snakes in air and in water with access to air. Frequency of air breathing, tidal volumes and total lung volumes were also compared in the two species. All measurements were done at 20–22 ° C.The aquaticAchrochordus showed long periods of apnea in submerged condition interrupted by short periods of breathing activity at the surface. Average frequency of air breathing activity was 2.6 times per hour. Breathing in constrictor was more frequent but irregular with an average frequency of 143 breaths per hour.Total lung volume was 66±31 ml/kg body weight and 72.5±59 ml/kg body weight inAchrochordus andConstrictor, respectively. Tidal volumes were 41.5±4.4 ml/kg body weight and 29.5±14.8 ml/kg body weight, largest inAchrochordus. Constrictor had the highest total O₂ uptake ( ) correlating with a higher activity. Total gas exchange ratio (R _E ) was 0.69 forConstrictor and 0.77 forAchrochordus. InConstrictor air breathing accounted for 97% of the total whereas 21% of the CO₂ exchange was aquatic. Corresponding figures forAchrochordus were 92% of total by air breathing with as much as 33% of the CO₂ elimination as aquatic gas exchange.The results demonstrate that the trend among early air breathing vertebrates (fishes and amphibians) of a conservative evolution of CO₂ elimination by air breathing also extends to snakes.Significantly the cutaneous exchange component was highest in the more aquatic species.The results are discussed in relation to recent reports of a higher than alleged role of the skin of reptiles in evaporative water loss.This study was supported by grant HE 12071 from the National Institutes of Health in the U. S. A.     

Breath-by-breath alveolar gas exchange

A method is described for breath-by-breath measurement of alveolar gas exchange corrected for changes of lung gas stores. In practice, the subject inspires from a spirometer, and each expired tidal volume is collected into a rubber bag placed inside a rigid box connected to the same spirometer. During the inspiration following any given expiration the bag is emptied by a vacuum pump. A computer monitors inspiratory and expiratory tidal volumes, drives four solenoid valves allowing appropriate operation of the system, and memorizes end-tidal gas fractions as well as mixed expired gas composition analyzed by mass spectrometer. Thus all variables for calculating alveolar gas exchange, based on the theory developed by Auchincloss et al. (J. Appl. Physiol. 21: 810-818, 1966), are obtained on a single-breath basis. Mean resting and steady-state exercise gas exchange data are equal to those obtained by conventional open-circuit measurements. Breathing rates up to 30 X min-1 can be followed. The breath-to-breath variability of O2 uptake at the alveolar level is less (25-35%) than that measured at the mouth as the difference between the inspired and expired volumes, both at rest and during exercise up to 0.7 of maximum O2 consumption.     

Diffusion in gas exchange of insects

The air-filled tracheal system constitutes the organ for gas exchange in terrestrial insects-its finest branches, the tracheoles, contacting individual cells. In the pupal stage, in which the animal lacks significant ventilatory movement, diffusion in the gas phase of the tracheal system constitutes the only mechanism for gas transfer between the environment and the tissues, transport in the hemolymph being insignificant. We have attempted to identify the main sites of diffusional resistance in the tracheal gas system by measuring the evolution of inert gases of low solubility from the pupa of the giant silkworm moth (Hyalophora cecropia). The results are compatible wih a single model in which the resistance to diffusional gas transfer in the tracheal system is concentrated at its opening at the body surface (spiracle).     

Pulmonary gas exchange in panting dogs

Pulmonary gas exchange during panting was studied in seven conscious dogs (32 kg mean body wt) provided with a chronic tracheostomy and an exteriorized carotid artery loop. The animals were acutely exposed to moderately elevated ambient temperature (27.5 degrees C, 65% relative humidity) for 2 h. O2 and CO2 in the tracheostomy tube were continuously monitored by mass spectrometry using a special sample-hold phase-locked sampling technique. PO2 and PCO2 were determined in blood samples obtained from the carotid artery. During the exposure to heat, central body temperature remained unchanged (38.6 +/- 0.6 degrees C) while all animals rapidly switched to steady shallow panting at frequencies close to the resonant frequency of the respiratory system. During panting, the following values were measured (means +/- SD): breathing frequency, 313 +/- 19 breaths/min; tidal volume, 167 +/- 21 ml; total ventilation, 52 +/- 9 l/min; effective alveolar ventilation, 5.5 +/- 1.3 l/min; PaO2, 106.2 +/- 5.9 Torr; PaCO2, 27.2 +/- 3.9 Torr; end-tidal-arterial PO2 difference [(PE' - Pa)O2], 26.0 +/- 5.3 Torr; and arterial-end-tidal PCO2 difference, [(Pa - PE')CO2], 14.9 +/- 2.5 Torr. On the basis of the classical ideal alveolar air approach, parallel dead-space ventilation accounted for 54% of alveolar ventilation and 66% of the (PE' - Pa)O2 difference. But the steepness of the CO2 and O2 expirogram plotted against expired volume suggested a contribution of series in homogeneity due to incomplete gas mixing.     

Nitrogen nutrition and water stress effects on leaf photosynthetic gas exchange and water use efficiency in winter wheat

The responses of gas exchange and water use efficiency to nitrogen nutrition for winter wheat were investigated under well-watered and drought conditions. The photosynthetic gas exchange parameters of winter wheat are remarkably improved by water and nitrogen nutrition and the regulative capability of nitrogen nutrition is influenced by water status. The effects of nitrogen nutrition on photosynthetic characteristics and on the limited factors to photosynthesis are not identical under different water status. Intrinsic water use efficiency (WUE(i)) of the plants at the high-N nutrition was decreased by a larger value than that of the plants in the low-N treatment due to a larger decrease in photosynthetic rate than in transpiration rate. Carbon isotope composition of plant material (delta(p)) is increased by the increase of drought intensity. The delta(p) at a given level of C(i)/C(a) is reduced by nitrogen deficiency. Leaf carbon isotope discrimination (Delta) is increased by the increase of nitrogen nutrition and decreased by the increase of drought intensity. Transpirational water use efficiency (WUE(t)) is negatively correlated with Delta in both nitrogen supply treatments and increased with the nitrogen supply.     

Leaf gas exchange in species of the Theobroma genus

Species of the Theobroma genus are primarily known by their commercially valuable seeds, especially, T. cacao is one of the most important tropical perennial crops. Beside T. grandiflorum, T. bicolor, and T. angustifolium, T. cacao is the only species of the genus that has been better studied to obtain physiologically relevant information. The main objective of this work was to evaluate the leaf gas exchange in seedlings of seven species of the Theobroma genus, seeking to identify characteristics that could be used in T. cacao breeding programmes. The study was realized under greenhouse conditions using six-month-old seedlings, in which net photosynthetic rate (P _N), stomatal conductance (g _s), transpiration (E), as well as parameters derived from light curves (P _N vs. photosynthetically active radiation) were evaluated. T. cacao, along with T. microcarpum, showed the lowest values of P _N, g _s, and E, while the highest values were presented by T. speciosum, which showed higher saturation irradiance and lower intrinsic and instantaneous water-use efficiencies, being considered the species less conservative in water use. Therefore, the parameters shown by the different evaluated species could serve to design T. cacao genotypes, through introgression of genes for specific environments such as the cabruca system widespread in southern Bahia, Brazil.     

Structural aspects of gas exchange

The lung is composed of several million small air spaces, lined by a delicate tissue membrane separating air from capillary blood. The design features of the gas exchange region in the lung are optimal for gaseous diffusion, by having a very extensive contact surface but with a minimal tissue barrier composed of an epithelial and endothelial layer separating an interstitial layer. The extent of the gas exchange surface in adult lungs is determined by general maturation which in turn is influenced by metabolic requirements of the organism. Environmental factors can modulate the pattern of ultimate lung development. Lung inflation causes air spaces to expand mainly by a process of tissue unfolding beneath an extremely thin layer of alveolar surfactant. This ensures cellular integrity during extreme deformations while at the same time providing a reserve of gas exchange surface so that functional diffusion capacity at all lung volumes is less than the structural maximum.     

Pulmonary gas exchange during exercise in pigs

Increased ventilation-perfusion(A/)inequality is observed in ~50% of humans during heavy exercise andcontributes to the widening of the alveolar-arterialO₂ difference(A-aD_O2). Despite extensive investigation, the cause remains unknown. As a firststep to more direct examination of this problem, we developed an animalmodel. Eight Yucatan miniswine were studied at rest and duringtreadmill exercise at ~30, 50, and 85% of maximalO₂ consumption (O_{2 max}). Multipleinert-gas, blood-gas, and metabolic data were obtained. TheA-aD_O2increased from 0 ± 3 (SE) Torr at rest to 14 ± 2 Torr duringthe heaviest exercise level, but arterialPO₂(Pa_O2) remained at resting levels during exercise. There was normalA/inequality [log SD of the perfusion distribution(log) = 0.42 ± 0.04] at rest, and moderate increases(log = 0.68 ± 0.04, P < 0.0001) wereobserved with exercise. This result was reproducible on a separate day.TheA/inequality changes are similar to those reported in highly trainedhumans. However, in swine, unlike in humans, there was no inert gasevidence for pulmonary end-capillary diffusion limitation during heavyexercise; there was no systematic difference in the measuredPa_O2 and the Pa_O2 as predicted from the inertgases. These data suggest that the pig animal model iswell suited for studying the mechanism of exercise-inducedA/ inequality.

     

Effects of human pregnancy and aerobic conditioning on alveolar gas exchange during exercise

This study examined the effects of aerobic conditioning during the second and third trimesters of human pregnancy on ventilatory responses to graded cycling. Previously sedentary pregnant women were assigned randomly to an exercise group (n = 14) or a nonexercising control group (n = 14). Data were collected at 15-17 weeks, 25-27 weeks and 34-36 weeks of pregnancy. Testing involved 20 W.min-1 increases in work rate to a heart rate of 170 beats.min-1 and (or) volitional fatigue. Breath-by-breath ventilatory and alveolar gas exchange measurements were compared at rest, a standard submaximal .VO2 and peak exercise. Within both groups, resting .V(E), .V(A), and V(T)/T(I) increased significantly with advancing gestation. Peak work rate, O2 pulse (.VO2/HR), .V(E), .V(A) respiratory rate, V(T)/T(I), .VO2, .VCO2, and the ventilatory threshold (T(vent)) were increased after physical conditioning. Chronic maternal exercise has no significant effect on pregnancy-induced changes in ventilation and (or) alveolar gas exchange at rest or during standard submaximal exercise. Training-induced increases in T(vent) and peak oxygen pulse support the efficacy of prenatal fitness programs to improve maternal work capacity.