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I R Cowan

Publications and source records attributed to I R Cowan.

9 recordsLinked to original sources

Gradients of Intercellular CO(2) Levels Across the Leaf Mesophyll.

Most current photosynthesis models, and interpretations of many wholeleaf CO(2) gas exchange measurements, are based on the often unstated assumption that the partial pressure of CO(2) is nearly uniform throughout the airspaces of the leaf mesophyll. Here we present measurements of CO(2) gradients across amphistomatous leaves allowed to assimilate CO(2) through only one surface, thus simulating hypostomatous leaves. We studied five species: Eucalyptus pauciflora Sieb. ex Spreng., Brassica chinensis L., Gossypium hirsutum L., Phaseolus vulgaris L., and Spinacia oleracea L. For Eucalyptus, maximum CO(2) pressure differences across the leaf mesophyll were 73 and 160 microbar when the pressures outside the lower leaf surface were 310 and 590 microbar, respectively. Using an approximate theoretical calculation, we infer that if the CO(2) had been supplied equally at both surfaces then the respective mean intercellular CO(2) pressures would have been roughly 12 and 27 microbar less than the pressures in the substomatal cavities in these cases. For ambient CO(2) pressures near 320 microbar, the average and minimum pressure differences across the mesophyll were 45 and 13 microbar. The corresponding mean intercellular CO(2) pressures would then be roughly 8 and 2 microbar less than those in the substomatal cavities. Pressure differences were generally smaller for the four agricultural species than for Eucalyptus, but they were nevertheless larger than previously reported values.

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Leaf Conductance in Relation to Rate of CO(2) Assimilation: I. Influence of Nitrogen Nutrition, Phosphorus Nutrition, Photon Flux Density, and Ambient Partial Pressure of CO(2) during Ontogeny.

Plants of Zea mays were grown with different concentrations of nitrate (0.6, 4, 12, and 24 millimolar) and phosphate (0.04, 0.13, 0.53, and 1.33 millimolar) supplied to the roots, photon flux densities (0.12, 0.5, and 2 millimoles per square meter per second), and ambient partial pressures of CO(2) (305 and 610 microbars). Differences in mineral nutrition and irradiance led to a large variation in rate of CO(2) assimilation per unit leaf area (A, 11 to 58 micromoles per square meter per second) when measured under standard conditions. The variation was shown, with the plants that had received different amounts of nitrate, to be related to variations in the nitrogen and chlorophyll contents, and phosphoenolpyruvate and ribulose-1,5-bisphosphate carboxylase activities per unit leaf area. Irrespective of growth treatment, A and leaf conductance to CO(2) transfer (g), measured under standard conditions were in almost constant proportion, implying that intercellular partial pressure of CO(2) (p(i)), was almost constant at 95 microbars. The same proportionality was maintained as A and g increased in an initially nitrogen-deficient plant that had been supplied with abundant nitrate. It was shown that p(i) measured at a given ambient partial pressure was not affected by the ambient partial pressure at which the plants had been grown, although it was different when measured at different ambient partial pressures. This suggests that the close coupling between A and g in these experiments is not associated with sensitivity of stomata to change in p(i).Similar, though less comprehensive, experiments were done with Gossypium hirsutum, and yielded similar conclusions, except that the proportionality between A and g at normal ambient partial pressure of CO(2) implied P(i) approximately 200 microbars.

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Leaf Conductance in Relation to Rate of CO(2) Assimilation: II. Effects of Short-Term Exposures to Different Photon Flux Densities.

When photon flux density incident on attached leaves of Zea mays L. was varied from the equivalent of 0.12 of full sunlight to full sunlight, leaf conductance to CO(2) transfer, g, changed in proportion to the change in rate of CO(2), assimilation, A, with the result that intercellular partial pressure of CO(2) remained almost constant. The proportionality was the same as that previously found in g and A measured at one photon flux density in plants of Zea mays L. grown at different levels of mineral nutrition, light intensities, and ambient partial pressures of CO(2). In shade-grown Phaseolus vulgaris L. plants, A as photon flux density was increased from about 0.12 up to about 0.5 full sunlight, the proportionality being almost the same in plants grown at low and at high light intensity.When photon flux density incident on the adaxial and abaxial surfaces of the isolateral leaves of Eucalyptus pauciflora Sieb. ex Spreng was varied, g and A also varied proportionally. The leaf conductance in a particular surface was affected by the photon flux density at the opposite surface to a greater extent than was expected on the basis of transmittance. The results indicated that stomata may, in some way, be sensitive to the photon flux absorbed within the leaf as a whole.

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Leaf Conductance in Relation to Rate of CO(2) Assimilation: III. Influences of Water Stress and Photoinhibition.

Rates of CO(2) assimilation and leaf conductances to CO(2) transfer were measured in plants of Zea mays during a period of 14 days in which the plants were not rewatered, and leaf water potential decreased from -0.5 to -8.0 bar. At any given ambient partial pressure of CO(2), water stress reduced rate of assimilation and leaf conductance similarly, so that intercellular partial pressure of CO(2) remained almost constant. At normal ambient partial pressure of CO(2), the intercellular partial pressure of CO(2) was estimated to be 95 microbars. This is the same as had been estimated in plants of Zea mays grown with various levels of nitrogen supply, phosphate supply and irradiance, and in plants of Zea mays examined at different irradiances.After leaves of Phaseolus vulgaris L. and Eucalyptus pauciflora Sieb. ex Spreng had been exposed to high irradiance in an atmosphere of CO(2)-free N(2) with 10 millibars O(2), rates of assimilation and leaf conductances measured in standard conditions had decreased in similar proportions, so that intercellular partial pressure of CO(2) remained almost unchanged. As the conductance of each epidermis that had not been directly irradiated had declined as much as that in the opposite, irradiated surface it was hypothesized that conductance may have been influenced by photoinhibition within the mesophyll tissue.

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A Direct Confirmation of the Standard Method of Estimating Intercellular Partial Pressure of CO(2).

The partial pressure of CO(2) inside leaves of several species was measured directly. Small gas exchange chambers were clamped above and below the same section of an amphistomatous leaf. A flowing gas stream through one chamber allowed normal CO(2) and water vapor exchange. The other chamber was in a closed circuit consisting of the chamber, an infrared gas analyzer, and a peristaltic pump. The CO(2) in the closed system rapidly reached a steady pressure which it is believed was identical to the CO(2) pressure inside the leaf, because there was no flux of CO(2) across the epidermis. This measured partial pressure was in close agreement with that estimated from a consideration of the fluxes of CO(2) and vapor at the other surface.

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Leaf Conductance in Relation to Assimilation in Eucalyptus pauciflora Sieb. ex Spreng: Influence of Irradiance and Partial Pressure of Carbon Dioxide.

Rates of assimilation and transpiration in Eucalyptus pauciflora Sieb. ex Spreng were measured at various ambient partial pressures of CO(2) and various irradiances and were used to estimate leaf conductance and intercellular partial pressure of CO(2). The responses of leaf conductance and rate of assimilation to change in intercellular partial pressure of CO(2) were expressed in terms of feedback. They are small in the sense that their combined effect was to reduce disturbances in intercellular partial pressure of CO(2) by 30% only. The magnitude of the feedback had no influence on the system as affected by irradiance, because the direct responses of conductance and rate of assimilation to changes in irradiance in the range 0.25 to 2 millieinsteins per meter per second were such that intercellular partial pressure was maintained almost constant.

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Oscillations in stomatal conductance: the influence of environmental gain.

It is supposed that oscillations in stomatal conductance are associated with the dynamic properties of the loop in which rate of evaporation affects, through physiological processes, the aperture of stomata and stomatal aperture in turn affects rate of evaporation. It is therefore predicted that their occurrence must be influenced by the magnitude of what is termed environmental gain: the sensitivity of rate of evaporation to change in leaf conductance to vapor transfer. Two methods of manipulating gain, and their effects on stomatal behavior in cotton (Gossypium hirsutum L. cv. Deltapine Smooth Leaf), are described. In the first, gain was increased by decreasing ambient humidity; in the second, it was made zero by regulating ambient humidity to keep rate of evaporation constant despite changes in conductance. The results are in accord with the supposition.

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Carbon dioxide exchange in cotton: some anomalous fluctuations.

Anomalous depressions in carbon dioxide exchange were observed in cotton leaves that were exhibiting oscillations in transpiration under controlled conditions of environment. The depressions occurred only when leaf temperature exceeded 37.5 degrees C and when the leaf diffusive resistance was minimum. Stomatal control of the supply of carbon dioxide to the leaf does not seem to be implicated in the effect.

Carbon Dioxide↗