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The lateral fluence-rate gradients in unilaterally irradiated maize (Zea mays L.) coleoptiles were calculated on the basis of the proportions of P fr (far-red-absorbing form of phytochrome) measured spectroscopically in transverse slices of the coleoptiles (top 1 cm). The results showed the occurrence of significant gradients that are wavelength-dependent. The gradient at 449 nm was steeper than those measured at 516, 534 and 551 nm, which were steeper than that measured at 665 nm. The ratios between the sides proximal and distal to the light source were, for example, 1:0.12 (449 nm), 1:0.23 (534 nm), and 1:0.28 (665 nm). Fluence-response curves for coleoptile phototropism (first positive curvature produced by less than 100 s unilateral irradiation) were measured at 449, 516, 534 and 551 nm. Comparison of the threshold fluences indicated that the responsiveness to 551 nm is about 10(4.8) less than that to 449 nm. Increasing wavelengths led to a decrease in maximal curvature, which correlated with the decrease of the fluence-rate ratios between the proximal and distal sides. Phototropic fluence-response curves were also measured using bilateral irradiation (449 nm). In one set of experiments, the fluence ratio was kept constant (either 1:1/2, 1:1/4 or 1:1/16) and the total fluence was varied, and in the other set the fluence applied to one side was kept constant and the fluence ratio was varied. A simple model based on the assumption that only one photoreaction occurs, and that the response is a function of the difference between the proximal and distal sides in the local photoreceptor action was tested. A fluence-response curve for this local photoreceptor action was calculated based on the fluence-rate ratio and the phototropic fluence-response curve measured for 449 nm. This curve was used, in conjunction with the measured fluence-rate ratios, as a basis for calculating phototropic fluence-response curves for other wavelengths and those for 449 nm obtained with bilateral irradiation. The calculated fluence-response curves showed excellent agreement with the experimental data. It is concluded that the threshold for maize coleoptile phototropism reflects the apparent photoconversion cross-section of the blue-light receptor whereas the maximal curvature depends on the steepness of the light gradient across the coleoptile.

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Unilateral irradiation of maize (Zea mays L.) seedlings results in a fluence-rate gradient, and hence below saturation, a gradient of the far-red-absorbing form of phytochrome (Pfr). The Pfr-gradients established by blue, red and far-red light were spectrophotometrically measured in the mesocotyl. Based on these Pfr-gradients and the fluence-response curves of phytochrome photoconversion the fluence-rate gradients were calculated. The fluence-rate gradient in the blue (460 nm) was steeper than that in the red (665 nm), which in turn was steeper than that in the far-red light (725 nm). The fluence-rate ratios front to rear were 1:0.06 (460 nm), 1:0.2 (665 nm), and 1:0.33 (725 nm). The assumption that phytochrome-mediated phototropism of maize mesocotyls is caused by local phytochrome-mediated growth inhibition was tested in the following manner. Firstly, the Pfr response curve for growth inhibition was calculated; these calculations were based on measurements of Pfr-gradients and data from red-light-induced phototropism. Secondly, the Pfr response curve for growth inhibition was used as a basis for calculating fluence-response curves for blue-and far-red-light-induced phototropism. Finally, these calculated results were compared with experimental data. It was concluded that the threshold for phytochrome-mediated phototropism of maize mesocotyls reflects the apparent photoconversion cross section of phytochrome whereas the maximal inducable curvature depends on the steepness of the light (Pfr) gradient across the mesocotyl.

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