Global-change scenarios suggest a trend of increasing diffuse light due to expected increases in cloud cover. Canopy-level measurements of plant-community photosynthesis under diffuse light show increased productivity attributed to more uniform distribution of light within the forest canopy, yet the effect of the directional quality of light at the leaf level is unknown. Here we show that leaflevel photosynthesis in sun leaves of both C3 and C4 plants can be 10-15% higher under direct light compared to equivalent absorbed irradiances of diffuse light. High-lightgrown leaves showed significant photosynthetic enhancement in direct light, while shade-adapted leaves showed no preference for direct or diffuse light at any irradiance. Highlight-grown leaves with multiple palisade layers may be adapted to better utilize direct than diffuse light, while shade leaf structure does not appear to discriminate light based on its directionality. Based upon our measurements, it appears that leaf-level and canopy-level photosynthetic processes react differently to the directionality of light, and previously observed increases in canopy-level photosynthesis occur even though leaf-level photosynthesis decreases under diffuse light.

In order to investigate the effects, without competition, of CO2 on germination, growth, physiological response, and reproduction, we focussed on co—occurring species that are prominent members of an annual community in Illinois. Five species of old field annual plants—Abutilon theophrasti (C3), Amaranthus retroflexus (C4), Ambrosia artemisiifolia (C3), Chenopodium album (C3), and Setaria faberii (C4)–were grown for their entire life cycle as individuals at CO2 concentration of 350 @mL/L, 500 @mL/L, and 700@mL/L. Emergence time, growth rate, shoot water status, photosynthesis, conductance, flowering time, nitrogen content, and biomass and reproductive biomass were measured. There was no detectable effect of enhanced CO2 on timing of emergence in any of the species. Amaranthus relative growth rate (RGR) was always higher at 700 @mL/L CO2 than at 350 @mL/L. In both Abutilon and Ambrosia, RGR was greater at 700 @mL/L than at 350 @mL/L during the first half of the experimental period, but during the second half of the period the reverse was true. Shoot water potential significantly increased (became less negative) with increasing CO2 in Amaranthus and Setaria. Similar but statistically nonsignificant trends were found in Chenopodium and Abutilon. Overall rate of photosynthesis increased with CO2 but there were no significant effects, at the species level, of CO2 on photosynthetic rates. Stomatal conductance decreased with increased CO2 at both high and low light levels in C3 species but only at high light levels in C4 species. In all species, intercellular CO2 increased with external CO2. Amaranthus flowered significantly earlier at 700 @mL/L than at 350 @mL/L, and Setaria flowered significantly later at 700 @mL/L than at either of the other CO2 levels. Both Abutilon and Ambrosia showed a trend towards earlier flowering but this was not statistically significant. Of the morphological characters measured at the final harvest only specific leaf area (SLA) showed a consistent response to CO2, decreasing with increasing CO2. Significant CO2 x species interactions were also found for leaf area, leaf biomass, biomass of reproductive parts, and seed biomass indicating species—specific responses for these characters. The proportion of nitrogen declined with increasing CO2: there was also a significant CO2 x species interaction caused by the different rates of decline in proportion of nitrogen among the species. The response of most characters had a significant species x CO2 interaction. However, this was not simply caused by the C3/C4 dichotomy. Reproductive biomass (seed, fruits, and flowers) increased with increasing CO2 in Amaranthus (C4) and in Chenopodium and Ambrosia (both C3), but there was no change in Setaria (C4), and Abutilon (C3) showed a peak a 500 @mL/L. Species of the same community differed in their response to CO2, and these differences may help explain the outcome of competitive interactions among these species above ambient CO2 levels.

Detailed growth analysis in conjunction with information on leaf display and nitrogen uptake was used to interpret competition between Abutilon theophrasti, a C annual, and Amaranthus retroflexus, a C annual, under ambient (350 μl l) and two levels of elevated (500 and 700 μl l) CO. Plants were grown both individually and in competition with each other. Competition caused a reduction in growth in both species, but for different reasons. In Abutilon, decreases in leaf area ratio (LAR) were responsible, whereas decreased unit leaf rate (ULR) was involved in the case of Amaranthus. Mean canopy height was lower in Amaranthus than Abutilon which may explain the low ULR of Amaranthus in competition. The decrease in LAR of Abutilon was associated with an increase in root/shoot ratio implying that Abutilon was limited by competition for below ground resources. The root/shoot ratio of Amaranthus actually decreased with competition, and Amaranthus had a much higher rate of nitrogen uptake per unit of root than did Abutilon. These latter results suggest that Amaranthus was better able to compete for below ground resources than Abutilon. Although the growth of both species was reduced by competition, generally speaking, the growth of Amaranthus was reduced to a greater extent than that of Abutilon. Regression analysis suggests that the success of Abutilon in competition was due to its larger starting capital (seed size) which gave it an early advantage over Amaranthus. Elevated CO had a positive effect upon biomass in Amaranthus, and to a lesser extent, Abutilon. These effects were limited to the early part of the experiment in the case of the individually grown plants, however. Only Amaranthus exhibited a significant increase in relative growth rate (RGR). In spite of the transitory effect of CO upon size in individually grown plants, level of CO did effect final biomass of competitively grown plants. Abutilon grown in competition with Amaranthus had a greater final biomass than Amaranthus at ambient CO levels, but this difference disappeared to a large extent at elevated CO. The high RGR of Amaranthus at elevated CO levels allowed it to overcome the difference in initial size between the two species.

We studied carbon dioxide concentrations in a mixed deciduous forest in New England, USA by making continuous measurements at 0.05, 0.2, 3, and 12 m above the soil surface. The measurements began in early March and continued until the end of November 1985; therefore, they spanned the growing season and parts of the dormant seasons both before and afterwards. The data were compared with those from Mauna Loa, Hawaii, which represent global atmospheric CO2 levels in the Northern Hemisphere. The results show strong seasonal and daily variations in CO2 concentrations at all heights in the forest. On average, during the growing season, CO2 levels were generally higher in the forest than in bulk air at Mauna Loa. The highest level of CO2 was found near the forest floor and the lowest at the 12—m level. Daily levels of CO2 were constant throughout the day in the dormant season and were the same for all heights in the forest. However, during the growing season, the CO2 concentrations were lowest during the middle of the day, especially at the 12 m height. Thus, this study shows that the CO2 concentrations in the forest may be quite different than those in bulk air and that seedlings, saplings, and mature trees may experience different CO2 environments for which they may show different responses in photosynthesis, growth, and water use. Moreover, a tree may experience different CO2 environments as it grows towards the canopy, and different modules of an individual may also be growing in different CO2 atmospheres.

Circadian rhythms in stomatal aperture and in stomatal conductance have been observed previously. Here we investigate circadian rhythms in apertures that persist in functionally isolated guard cells in epidermal peels of Vicia faba, and we compare these rhythms with rhythms in stomatal conductance in attached leaves. Functionally isolated guard cells kept in constant light display a rhythmic change in aperture superimposed on a continuous opening trend. The rhythm free-runs with a period of about 22 hours and is temperature compensated between 20 and 300C. Functionally isolated guard cell pairs are therefore capable of sustaining a true circadian rhythm without interaction with mesophyll cells. Stomatal conductance in whole leaves displays a more robust rhythm, also temperature-compensated, and with a period similar to that observed for the rhythm in stomatal aperture in epidermal peels. When analyzed individually, some stomata in epidermal peels showed a robust rhythm for several days while others showed little rhythmicity or damped out rapidly. Rhythmic periods may vary between individual stomata, and this may lead to desynchronization within the population. Stomata function to control CO2 uptake and transpirational water loss, and they are closely regulated by such factors as light (for review see 23, 28), RH (for review see 22), and carbon dioxide concentration (for review see 17). In addition to environmental factors, stomata are controlled by an endogenous circadian clock. This control may appear as a rhythmic change in aperture under constant conditions (15,21) or as a rhythmic change in sensitivity to some environmental factor; light has been studied most frequently (4, 5, 15). Here we are concerned only with the first type of endogenous control: simple, free-running rhythmicity in stomatal aperture. A wide variety of plant species show such free-

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We review the influence of self-focusing on the measurement of bulk laser-induced-damage (LID) thresholds in normally transparent optical materials. This role is experimentally determined by measuring the spot size and polarization dependence of LID and by observing beam distortion in the far field. Utilizing these techniques, we find that by using a tight focusing geometry in which the breakdown power is below P2, the effects of self-focusing can be practically eliminated in an LID experiment. P2 is the so-called second critical power for self-focusing, and P2 = 3.77P1, where P1 = cÀ2 /3272n2, where c is the speed of light in vacuum, X is the laser wavelength and n2 is the nonlinear index of refraction. This is in accordance with numerical calculations by J. H. Marburger [in Progress in Quantum Electronics, J. H. Sanders and S. Stenholm, eds., Vol. 4, Part 1, pp. 35-110, Pergamon, Oxford (1975)]. With this knowledge we determine that damage is only partially explained by avalanche ionization and that the initiation of damage is strongly influenced by extrinsic processes. We therefore conclude that we are measuring extrinsic LID.