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Abstract. Because CO2 diffuses 10000 times more slowly through water than air, there may be strong selective pressure for increased water repellency in terrestrial plant leaves. In the present study, leaf trichomes appeared to have a strong influence on leaf water repellency (i.e. degree of water droplet formation on the leaf surface) as well as the retention of droplets on the leaf. Based upon evaluation of 38 plant species from 21 families, we found that leaves with trichomes were more water repellent, especially where trichome density was greater than 25mm2. However, droplet repellency and retention were both high in some species where trichomes entrapped droplets. Finally, the lensing effects of water droplets on leaf surfaces increased incident sunlight by over 20‐fold directly beneath individual droplets. These results may have important implications for such processes as stomatal function, whole leaf photosynthesis, and transpiration for a large variety of plant species.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. University of California Press and American Institute of Biological Sciences are collaborating with JSTOR to digitize, preserve and extend access to BioScience. Morphological and anatomical features of plant leaves are commonly associated with metabolic type (e.g., Kranz anatomy of C4 species), amount of sun exposure (e.g., sun and shade leaves), or water stress (e.g., xeromorphism). However, although the primary function of the leaf is to absorb and process sunlight and carbon dioxide for photosynthesis, few structural features of leaves have been related mechanistically to these tasks. For example, it has been known for over a century that the internal anatomy of leaves is characterized by different cell layers (e.g., the palisade and spongy mesophyll) and that stomatal pores can be located on one or both sides of a leaf. Yet, only recently has any functional relationship between leaf form and photosynthetic performance been suggested.A variety of ecological studies have correlated numerous leaf structural parameters with photosynthetic performance (e.g., Abrams and Kubiske 1990, 1994, Hinckley et al. 1989, William K. Smith (e-mail: wksmith@uwyo. edu) and Thomas C. Vogelmann (e-mail: tvogel@uwyo.edu) are professors in the

Light gradients were measured in leaves that had different types of anatomical development of the mesophyll but similar pigment content. Leaves of the legume, Thermopsis montana, had columnar palisade and spongy mesophyll whereas leaves of the monocot, Smilacina stellata, had spongy mesophyll only. Light gradients were measured at 550 nm in both types of leaves when they were irradiated with collimated or diffuse light. When irradiated with collimated light, light gradients were steeper in leaves with spongy mesophyll in comparison to those that had palisade tissue. On the other hand, light gradients were similar between both leaf types when they were irradiated with difTuse light. Thus, columnar palisade cells facilitated the penetration of collimated light over diffuse light. These results suggest that palisade tissue may help distribute light more uniformly to chloroplasts within the leaf. Moreover, the functional significance of palisade tissue may be related to the amount of collimated light within the natural environment.

Maximal l4 CO 2-fixation in spinach occurs in the middle of the palisade mesophyll [Nishio et al. (1993) Plant Cell 5: 953], however, ninety percent of the blue and red light is attenuated in the upper twenty percent of a spinach leaf [Cui et al. (1991) Plant Cell Environ. 14: 493]. In this report, we showed that green light drives 14 C0 2-fixation deep within spinach leaves compared to red and blue light. Blue light caused fixation mainly in the palisade mesophyll of the leaf, whereas red light drove fixation slightly deeper into the leaf than did blue light. I4 C0 2-fixation measured under green light resulted in less fixation in the upper epidermal layer (guard cells) and upper most palisade mesophyll compared to red and blue light, but led to more fixation deeper in the leaf than that caused by either red or blue light. Saturating white, red, or green light resulted in similar maximal 14 CO 2-fixation rates, whereas under the highest irradiance of blue light given, carbon fixation was not saturated, but it asymptotically approached the maximal 14 CO 2-fixation rates attained under the other types of light. The importance of green light in photosynthesis is discussed.

Abstract. Light gradients were measured and correlated with chlorophyll concentration and anatomy of leaves in spinach (Spinacia oleracea L.). Light gradients were measured at 450, 550 and 680 nm within thin (455 μm) and thick (630 μm) leaves of spinach grown under sun and shade conditions. The light gradients were relatively steep in both types of leaves and 90% of the light at 450 and 680 nm was absorbed by the initial 140 μm of the palisade. In general, blue light was depleted faster than red light which, in turn was depleted faster than green light. Light penetrated further into the thicker palisade of sun leaves in comparison to the shade leaves. The distance that blue light at 450 nm travelled before it became 90% depleted was 120 μm in sun leaves versus 76 μm in shade leaves. Red light at 680 nm and green light at 550 nm travelled further but the trends were similar to that measured at 450nm. The steeper light gradients within the palisade‐of shade leaves were caused by increased scattering of light within the intercellular air spaces and/or cells which were less compact than those in sun leaves. The decline in the amount of light within the leaf appeared to be balanced by a gradient in chlorophyll concentration measured in paradermal sections. Progressing from the adaxial epidermis, chlorophyll content increased through the palisade and then declined through the spongy mesophyll. Chlorophyll content was similar in the palisade of both sun and shade leaves. Chloroplast distribution within both sun and shade leaves was relatively uniform so that the chlorophyll gradient appeared to be caused by greater amounts of chlorophyll within chloroplasts located deeper within the leaf. These results indicate that the anatomy of the palisade may be of special importance for controlling the penetration of photo‐synthetically active radiation into the leaf. Changing the structural characteristics of individual palisade cells or their arrangement may be an adaptation that maximizes the absorption of light in leaves with varying mesophyll thickness due to different ambient light regimes.

Chlorophyll fluorescence was used to estimate profiles of absorbed light within chlorophyll solutions and leaves. For chlorophyll solutions, the intensity of the emitted fluorescence declined in a log-linear manner with the distance from the irradiated surface as predicted by Beer's law. The amount of fluorescence was proportional to chlorophyll concentration for chlorophyll solutions given epi-illumination on a microscope slide. These relationships appeared to hold for more optically complex spinach leaves. The profile of chlorophyll fluorescence emitted by leaf cross sections given epi-illumination corresponded to chlorophyll content measured in extracts of leaf paradermal sections. Thus epifluorescence was used to estimate relative chlorophyll content through leaf tissues. Fluorescence profiles across leaves depended on wavelength and orientation, reaching a peak at 50-70 µ µ µ µ m depth. By infiltrating leaves with water, the pathlengthening due to scattering at the airspace : cell wall interfaces was calculated. Surprisingly, the palisade and spongy mesophyll had similar values for pathlengthening with the value being greatest for green light (550 > 650 > 450 nm). By combining fluorescence profiles with chlorophyll distribution across the leaf, the profile of the apparent extinction coefficient was calculated. The light profiles within spinach leaves could be well approximated by an apparent extinction coefficient and the Beer-Lambert/ Bouguer laws. Light was absorbed at greater depths than predicted from fibre optic measurements, with 50% of blue and green light reaching 125 and 240 µ µ µ µ m deep, respectively.

Leaf anatomy plays a functional role in propagating light through the leaf; palisade mesophyll has been shown to facilitate the channelling of collimated light deeper into the spongy mesophyll. Direct measurements of the propagation of diffuse light into the leaf, however, are absent. Using chlorophyll fluorescence imaging of leaf cross-sections, we measured light absorption profiles in leaves under direct (collimated), diffuse and low-angle monochromatic light. Low-angle and diffuse light was absorbed closer to the irradiated surface than direct light perpendicular to the surface. The shapes of internal absorption profiles indicated that leaves were influenced by the directional quality of the incident light. In addition, absorption profiles revealed that leaves were not simple light absorbing objects and that cellular anatomy influences the direction of light travelling into the mesophyll. These findings also suggest a mechanism for previously measured differences in leaf level photosynthesis under opposing light regimes.