Further Studies on Differentiation of Photosynthetic Properties in Sun and Shade Ecotypes of Solidago virgaurea

Measurements of the fraction of the incident light absorbed by diverse Solidago leaves revealed that differences in light harvesting capacity cannot explain the differences in efficiency of utilization of weak light in photosynthesis that have previously been shown to exist between sun and shade ecotypes when these have been grown in strong light and between identical clones of shade ecotypes when grown at different light intensities. Photosynthesis measurements at low and normal oxygen concentrations, provided no evidence that a different degree of inhibition of photo-synthetic CO₂ uptake by atmospheric oxygen is responsible for the observed differences in photosynthetic efficiency, at low or high light intensities. These results support the conclusion that the markedly less efficient use of weak light by shaded habitat clones grown in strong as compared with weak light is caused primarily by damage to the photosystems, or to a site close to them. Measurements of Emerson enhancement and of light-induced absorbance changes provide some evidence that photoreaction II is more affected than I. Enzyme extracts prepared from clones native to an exposed habitat were found to contain considerably higher activities of carboxydismutase (ribulosc-l,5-diphos-phate carboxylase) than from clones native to a shaded habitat when the plants were previously grown at a moderately high light intensity. Exposed habitat clones apparently have a genetically determined, higher capacity to produce the carboxyla-tion enzyme than shaded habitat clones. The high degree of correlation found when the light-saturated rate of CO₂ uptake in vivo of a number of individual Solidago leaves is plotted against the carboxydismutase activities found in the extracts of these same leaves suggests that low carboxydismutase activity is one of the intrinsic properties responsible for the low capacity for light-saturated photosynthesis of clones from shaded habitats. It is concluded from this and other investigations that differentiation between plants from habitats with contrasting light intensities, whether unrelated species or ecotypos of the same species, probably involves the capacity of several component steps of the photosynthetic process.     

Photosynthetic response to varying light intensity in ecotypes of Solanum dulcamara L. from shaded and exposed habitats

Summary Within the widespread species Solanum dulcamara, contrasting ecotypes were found which are physiologically adapted to the light intensities prevailing in their natural habitats. When grown under a high light intensity, an ecotype from a shaded habitat exhibits signs of damage. Another one from an exposed habitat has higher rates of photosynthetic CO₂ uptake when grown under strong as compared to weak light and does not show damage. This differential response becomes even more evident when leaves of both ecotypes are grown to maturity under weak light and are subsequently subjected to strong light for some time. The quantum requirement for photosynthesis increases in the shade-, but not in the sun-ecotype. The sun type increases its rate of photosynthesis under saturating light intensities after a few days in strong light.No significant difference in physical resistances to gas diffusion could be found to explain the highly differing rates of photosynthesis. With the increase in photosynthetic capacity in leaves of the sun type, protein content, activity of RuDP carboxylase, and concentration of Fraction I protein increased likewise. It is suggested that de novo synthesis of photosynthetic enzymes in fully expanded leaves of the sun ecotype following treatment with strong light is the cause of its increased capacity for CO₂ fixation.     

Sun/shade adaptations of the photosynthetic apparatus of <Emphasis Type="Italic">Hoya carnosa</Emphasis>, an epiphytic CAM vine,in a subtropical rain forest in northeastern Taiwan

Most past work on the ecophysiology of the Crassulacean acid metabolism (CAM) plant, Hoya carnosa, in the lab and in situ in Australia indicates that this epiphytic vine is better adapted to shaded, not exposed, locations, although a recent study of this species in Taiwan presents findings that run counter to this conclusion. Thus, photosynthetic characteristics of shaded and exposed individuals of H. carnosa were compared in situ in a subtropical rain forest in northeastern Taiwan in order to determine whether this CAM epiphyte is better adapted to the shade or the sun. Although leaves of shade plants had much greater chlorophyll concentrations than did those of sun plants, chlorophyll a/b ratios did not differ between the two groups of plants. Fluorescence measurements revealed some ability of leaves to acclimate to both shade and sun, although some evidence for photoinhibition (photoprotection) was observed in more exposed plants. Despite the latter, both exposed and shaded plants exhibited CAM, measured as diel fluctuations in leaf acidity, and CAM was more consistently found in the exposed plants. Furthermore, some evidence for more CAM at higher light availabilities was found. Overall, the results of this investigation reveal that H. carnosa in this subtropical rain forest in Taiwan exhibits adaptations to both high and low light levels, which should prove adaptive for an epiphytic vine with leaves on the same individual exposed to a wide range of exposure and shade in the host tree canopy.     

SUN-SHADE ECOTYPES OF A BLUEGREEN ALGA IN A HOT SPRING1,2

Clone cultures of the thermophilic alga Plectonema notatum Schmidle were established from cells collected from the high and low light intensity regions of the algal mat which developed in Jerry Johnson Hot Spring, Idaho. Clones isolated from cither high, or low light intensify zones were grown at light intensities of 8000 and 400 ft-c. The existence of specialized and genetically fixed sun or shade ecotypes was evidenced, by the ability of low light intensity clones to synthesize more light harvesting chlorophyll a when grown under low light conditions than the high light clones. High light clones showed light saturation of photosynthesis at higher light intensities with higher carboxylating enzyme activities and less chlorophyll a than low light clones when both were cultured at 8000 ft-c. These clones displayed infraspecific variation along the light intensity gradient and therefore exist as a mosaic of light, intensity ecotypes.     

Adaptation of the Photosynthetic Apparatus of Scenedesmus obliquus to Strong and Weak Light Conditions

Homocontinuous cultures of the unicellular green alga Scenedesmus obliquus were grown under strong (28 W/m²～28,000 lux) and weak (5 W/m²～5000 lux) light conditions to simulate the conditions of ‘sun’ and ‘shade’ plants. As in higher plants the cells adapted to strong light had less chlorophyll but demonstrated a higher photosynthetic capacity and a higher respiration rate, so that their compensation point was reached at three times higher energy than in the cells grown under low light intensities. The CO₂ fixation rate and the RuDP carboxylase activity under saturating light intensities were both higher in the cells grown in strong light. In spite of the differences in the pigment content and in the light saturated photosynthetic capacities for both cultures, the quantum yields of photosynthetic oxygen evolution were equal. As documented for some species of higher plants Scenedesmus is not genetically determined to be either a ‘sun’ or ‘shade’ organism but can adapt its photosynthetic apparatus to the different light intensities.     

Photosynthetic adaptation of Solanum dulcamara L. to sun and shade environments

Summary The photosynthetic response properties of individuals of Solanum dulcamara L. collected from sun and shade habitats were compared in controlled environments. Light-saturated photosynthetic rates and seven additional parameters associated with photosynthetic and growth performance were measured over a range of 12 environmental conditions that simulated natural habitat differences in light intensity, moisture availability and daily temperature amplitude. In contrast to previous studies, the results suggest there is no ecotypic differentiation with respect to the sun and shade environments from which the individuals were collected. It appears that all but one of the field-collected individuals are capable of successfully inhabiting the full range of light environments from which the species was collected.     

Sun-shade adaptability of the red mangrove,Rhizophora mangle (Rhizophoraceae): Changes through ontogeny at several levels of biological organization

Rhizophora mangle L., the predominant neotropical mangrove species, occupies a gradient from low intertidal swamp margins with high insolation, to shaded sites at highest high water. Across a light gradient, R. mangle shows properties of both “light-demanding” and “shade-tolerant” species, and defies designation according to existing successional paradigms for rain forest trees. The mode and magnitude of its adaptability to light also change through ontogeny as it grows into the canopy. We characterized and compared phenotypic flexibility of R. mangle seedlings, saplings, and tree modules across changing light environments, from the level of leaf anatomy and photosynthesis, through stem and whole-plant architecture. We also examined growth and mortality differences among sun and shade populations of seedlings over 3 yr. Sun and shade seedling populations diverged in terms of four of six leaf anatomy traits (relative thickness of tissue layers and stomatal density), as well as leaf size and shape, specific leaf area (SLA), leaf internode distances, disparity in blade–petiole angles, canopy spread: height ratios, standing leaf numbers, summer (July) photosynthetic light curve shapes, and growth rates. Saplings showed significant sun/shade differences in fewer characters: leaf thickness, SLA, leaf overlap, disparity in bladepetiole angles, standing leaf numbers, stem volume and branching angle (first-order branches only), and summer photosynthesis. In trees, leaf anatomy was insensitive to light environment, but leaf length, width, and SLA, disparities in bladepetiole angles, and summer maximal photosynthetic rates varied among sun and shade leaf populations. Seedling and sapling photosynthetic rates were significantly depressed in winter (December), while photosynthetic rates in tree leaves did not differ in winter and summer. Seasonal and ontogenetic changes in response to light environment are apparent at several levels of biological organization in R. mangle, within constraints of its architectural baiiplan. Such variation has implications for models of stand carbon gain, and suggest that response flexibility may change with plant age.     

Drought-Responsive Gene Expression in Sun and Shade Plants of <Emphasis Type="Italic">Haberlea rhodopensis</Emphasis> Under Controlled Environment

Haberlea rhodopensis is a homoiochlorophyllous desiccation-tolerant plant growing mostly in shaded rock rifts below the trees at very low light intensity. These shade plants are very sensitive to photoinhibition and do not survive desiccation at irradiance of 350 μmol m^?2 s^?1, whereas plants growing on the top of rocks exposed to full sunlight (sun plants) can survive at even higher light intensities regularly. The aim of the present study was to establish how acclimation to different light intensities influences the expression of selected drought-responsive genes and the physiological activity during desiccation of shade and sun plants under controlled culture conditions. The photosynthetic activity was higher in sun plants not only when fully hydrated but also during dehydration. Thus, the higher photosynthetic capacity, reflected in PSII but especially in PSI activity, is accompanied by a reduced susceptibility to photodamage. For most of the genes examined, drought was the main factor in regulation; in addition, some were light modulated like genes coding for putative superoxide dismutase (SOD), ascorbate peroxidase (APX) and thioredoxin (TRX), whereby the former was almost purely light regulated. Differences between sun and shade plants concerned mainly on the time course. Whereas some genes reacted already at moderate desiccation only in sun plants (genes for monodehydroascorbate reductase (MDAR), plastidic translocase (PTL) similar to OEP16 and one of the genes, newly annotated ELIP-like, specific for H. rhodopensis), especially a gene for a putative UDP-glucuronic acid decarboxylase (UDP) retained its enhanced expression longer during recovery. Thus, these genes are probably especially important for survival and recovery in sun plants.     

Characterization of pigment apparatus in winter-green and summer-green leaves of a shade-tolerant plant <Emphasis Type="Italic">Ajuga reptans</Emphasis>

Comparative study was performed to assess the content and proportions of photosynthetic pigments and the violaxanthin cycle (VXC) activity in winter-green and summer-green leaves of bugleweed (Ajuga reptans L.) plants grown in shaded (photosynthetically active radiation, PAR 150 μmol/(m² s)) and sunny (PAR 1200 μmol/(m² s)) habitats in the Botanic Garden of Jagiellonian University (Krakow, Poland). In overwintered and newly formed leaves of shade plants, the content of green and yellow pigments was two times higher than in leaves of sun plants. The shade plants were distinguished by accumulation of β-carotene, while lutein was predominant in leaves of sun plants. Under the action of strong light (2000 μmol/(m²s)), the level of violaxanthin deepoxidation in winter-green leaves of shade and sun plants increased five- to sixfold, whereas it changed insignificantly in summer-green leaves of shade plants. It is concluded that, in a shadetolerant species A. reptans, the photosynthetic apparatus of winter-green leaves in sun and shade plants and of summer-green leaves in sun plants is protected against excess insolation by high activity of VXC. The carotenoids of summer-green leaves in shade plants are supposed to function mainly as light-harvesting pigments.     

Photosynthetic characteristics and chloroplast ultrastructure of C3 and C4 tree species grown in high- and low-light environments

Plants of the C₄ tree species, Euphorbia forbesii, Sherff and the C₃ tree species, Claoxylon sandwicense Muell-Arg., were grown in a full sun and a shade environment designed to simulate the understory of their native Hawiian forest habitat. When grown under shade conditions, both species exhibited a photosynthetic light response typical of shade plants with low light compensation points and low dark respiration rates. E. forbesii, however, exhibited greater acclimation of light saturated photosynthetic rates and no evidence of photoinhibition in high light. In contrast, quantum yields for CO₂ uptake and chlorophyll contents were reduced in the high-light as compared to the low-light grown C. sandwicense plants. Both species exhibited similar changes in the intercellular CO₂ response curves and chloroplast whole-chain electron transport capacities, suggesting that the underlying mechanisms of light acclimation are similar. Chloroplasts of E. forbesii exhibited large changes in ultrastructure, with much greater thylakoid membrane development in low than high light. In contrast, C. sandwicense exhibited different starch contents, but otherwise similar membrane development in high and low light. The results show that E. forbesii possesses a very flexible photosynthetic apparatus which may account for its ability to survive in the understory of shaded forests.Abbreviations g_s = stomatal conductance - HL = high light - LL = low light - P_i = intercellular CO₂ partial pressure - PFD = photon flux density     

Effect of light intensity and temperature on apparent photosynthesis of altitudinal ecotypes of Trifolium repens L.

Summary The dependence on light and temperature of the apparent photosynthetic rate was studied on ecotypes of Trifolium repens from different altitudes in the alps (600–2040 m above sea level). Due to the altitude, the natural habitats have different temperature conditions. At the higher altitudes the light conditions for the growing plants vary due to grazing or cutting management of these meadows. Accordingly, for this study the plants were grown at different temperatures and light intensities in growth cabinets.High altitude plants had higher photosynthetic rates, especially when measured at low temperatures. According to the light conditions, dependent on management, in the alpine habitats the ecotypes differed in their photosynthetic properties like sun and shade plants. It is stated, that the photosynthetic performance as well as the acclimation capacity to the growth conditions is related to the altitude of the habitats and probably also to the agricultural management.     

Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Plants of Solidago virgaurea L. from exposed and shaded habitats differ with respect to the response of the photosynthetic apparatus to the level of irradiance during growth. An analysis was carried out on leaf characteristies which might be responsible for the differences established in the rates of Hght-saturated CO₂ uptake. The clones were grown in controlled environment chambers at high and low levels of irradiance. Light-saturated rates of photosynthesis and transpiration were measured at natural and lower ambient CO₂ concentrations. A low temperature dependence of light-saturated CO₂ uptake at natural CO₂ concentrations, and a strong response to changes in stomatal width, suggested that the rate of CO₂ transfer from ambient air towards reaetion sites in chloroplasts was mainly limiting the pholosynthetic rate. Resistances to transfer of CO₂ for different parts of the pathway were calculated. There was a weak but significant correlation between stomatal conductance and the product stomatal frequency ± pore length. Mesopbyll conductance and dry weight per unit area were highly correlated in leaves not damaged by high irradiance. This suggests that mesophyll conductance increases with increasing cross sectional area (per unit leaf area) of the pathways of CO₂ transfer in the mesophyll from cell surfaces to reaction sites. The higher light-saturated photosynthesis in clones from exposed habitats when grown at high irradiance than when grown at low irradiance was attributable mainly to a lower mesophyll resistance. In shade clones the effect upon CO₂ uptake of the increase in leaf thickness when grown at high irradiance was counteracted by the associated inactivation of the photosynthetic apparatus. The difference in CO₂ uptake present between clones from exposed and shaded habitats when preconditioned to high irradiance resulted from differences in both mesophyll and stomatal resistances. A few hybrid clones of an F₁-population from a cross between a clone from an exposed habitat and a clone from a shaded habitat reacted, on the whole, in the same way as the exposed habitat parent. When grown at high irradiance, the hybrid clones showed higher photosynthetic rates than either parent; this was largely attributable to the unusually low stomatal resistance of the hybrid leaves.     

Acclimation of photosynthesis to lightflecks in tomato leaves: interaction with progressive shading in a growing canopy

Plants in natural environments are often exposed to fluctuations in light intensity, and leaf‐level acclimation to light may be affected by those fluctuations. Concurrently, leaves acclimated to a given light climate can become progressively shaded as new leaves emerge and grow above them. Acclimation to shade alters characteristics such as photosynthetic capacity. To investigate the interaction of fluctuating light and progressive shading, we exposed three‐week old tomato (Solanum lycopersicum ) plants to either lightflecks or constant light intensities. Lightflecks of 20 s length and 1000 μmol m^?2 s^?1 peak intensity were applied every 5 min for 16 h per day, for 3 weeks. Lightfleck and constant light treatments received identical daily light sums (15.2 mol m^?2 day^?1). Photosynthesis was monitored in leaves 2 and 4 (counting from the bottom) during canopy development throughout the experiment. Several dynamic and steady‐state characteristics of photosynthesis became enhanced by fluctuating light when leaves were partially shaded by the upper canopy, but much less so when they were fully exposed to lightflecks. This was the case for CO₂‐saturated photosynthesis rates in leaves 2 and 4 growing under lightflecks 14 days into the treatment period. Also, leaf 2 of plants in the lightfleck treatment showed significantly faster rates of photosynthetic induction when exposed to a stepwise change in light intensity on day 15. As the plants grew larger and these leaves became increasingly shaded, acclimation of leaf‐level photosynthesis to lightflecks disappeared. These results highlight continuous acclimation of leaf photosynthesis to changing light conditions inside developing canopies.     

Photosynthetic responses of C3 and C4 species from cool shaded habitats in Hawaii

Summary The C₄ species, Euphorbia forbesii, and the C₃ species, Claoxylon sandwicense, occupy cool, shaded habitats in Hawaii. Both of these species exhibit the photosynthetic characteristics of typical shade plants: low light-saturated photosynthetic rates, low dark respiration rates, low light levels for saturation of photosynthesis, and low light compensation points. In addition, the quantum yields of the two species are similar at leaf temperatures near 22°C, reflecting a significant increase in the quantum yield of E. forbesii over that of C₄ species from open habitats. C. sandwicense has a lower dark respiration rate than E. forbesii. Hence, since the quantum yields of the two species are similar at cool temperatures, C. sandwicense has a higher photosynthetic rate than E. forbesii at low incident photon flux densities. As a consequence, C. sandwicense should have a greater carbon gain than E. forbesii under the diffuse radiation conditions of their native habitat. However, since E. forbesii has a higher light-saturated photosynthetic rate than C. sandwicense, E. forbesii may have a greater carbon gain than C. sandwicense during sunflecks.     

Growth, biomass allocation and photosynthesis of invasive and native Hawaiian rainforest species

Growth, biomass allocation, and photosynthetic characteristics of seedlings of five invasive non-indigenous and four native species grown under different light regimes were studied to help explain the success of invasive species in Hawaiian rainforests. Plants were grown under three greenhouse light levels representative of those found in the center and edge of gaps and in the understory of Hawaiian rainforests, and under an additional treatment with unaltered shade. Relative growth rates (RGRs) of invasive species grown in sun and partial shade were significantly higher than those for native species, averaging 0.25 and 0.17 g g⁻¹ week⁻¹, respectively, while native species averaged only 0.09 and 0.06 g g⁻¹ week⁻¹, respectively. The RGR of invasive species under the shade treatment was 40% higher than that of native species. Leaf area ratios (LARs) of sun and partial-shade-grown invasive and native species were similar but the LAR of invasive species in the shade was, on average, 20% higher than that of native species. There were no differences between invasive and native species in biomass allocation to shoots and roots, or in leaf mass per area across light environments. Light-saturated photosynthetic rates (Pmax) were higher for invasive species than for native species in all light treatments. Pmax of invasive species grown in the sun treatment, for example, ranged from 5.5 to 11.9 μmol m⁻² s⁻¹ as compared with 3.0−4.5 μmol m⁻² s⁻¹ for native species grown under similar light conditions. The slope of the linear relationship between Pmax and dark respiration was steeper for invasive than for native species, indicating that invasive species assimilate more CO₂ at a lower respiratory cost than native species. These results suggest that the invasive species may have higher growth rates than the native species as a consequence of higher photosynthetic capacities under sun and partial shade, lower dark respiration under all light treatments, and higher LARs when growing under shade conditions. Overall, invasive species appear to be better suited than native species to capturing and utilizing light resources, particularly in high-light environments such as those characterized by relatively high levels of disturbance. Received: 30 December 1997 / Accepted: 1 September 1998     

Variations in leaf morpho‐anatomy and photosynthetic traits between sun and shade populations of Eurya japonica (Pentaphylacaceae) whose seeds are dispersed by birds across habitats

Eurya japonica occurs in diverse light environments through seed dispersal by birds. As the seed size is extremely small, we hypothesized that newly germinated seedlings with restricted depth of roots and length of the hypocotyl would suffer high mortality due to increased transpiration in sunny habitats and low light in shady habitats. We also expected that surviving seedlings would differ in leaf traits between habitats as a result of selection. We aimed to determine how photosynthetic traits differ between habitats and how leaf structure is related to this difference. We examined photosynthesis and leaf morpho‐anatomy for plants cloned from cuttings collected from the forest understory (shade population) and neighboring roadsides and cut‐over areas (sun population) and then grown under two irradiances (18.5% and 100% sunlight) in an experimental garden. Under growth in 100% sunlight, cloned plants from the sun population exhibited significantly greater area‐based photosynthetic capacity compared to cloned plants from the shade population at a comparable stomatal conductance, which was attributable to a higher area‐based leaf nitrogen concentration. On the other hand, mean values of photosynthetic capacity did not significantly differ between the two populations. Cloned plants from the sun population had significantly thicker leaf laminas and spongy tissue and lower stomatal density compared to cloned plants from the shade population. Thickened leaf lamina might have increased leaf tolerance to physical stresses in open habitats. The variation in leaf morpho‐anatomy between the two populations can be explained in terms of the economy of leaf photosynthetic tissue.     

Morphological plasticity and ecophysiological response of ground ivy (Glechoma hederacea,Lamiaceae) in contrasting natural habitats within its native range

Abstract

Some species intrinsically have a high invasiveness capacity, shown by high phenotypic plasticity and rapid growth, enabling a wide distribution across their native habitats and successful invasion in the introduced range. For such species, information from native habitats is critically important. An example is Glechoma hederacea, native to Eurasia but introduced and widespread in the USA. Our main objective was to investigate variation in traits of native G. hederacea populations across contrasting habitats: open, forest edge and understory. Vegetation was sampled and the ecophysiological and morphological traits were measured with accompanying environmental parameters. Results showed that in native habitats environmental conditions cover wide gradients of light and soil moisture. Plants had the highest cover in nutrient-rich, shaded habitats, representing the optimal habitat, indicating shade tolerance of G. hederacea. Plants from forest understory exhibited strong similarities in investigated traits to plants from the forest edge, even though this was a drier, sunnier habitat. Plants from open, sunny habitats experienced stress as indicated by the quantum efficiency of PSII and significantly higher sexual reproduction. Results show that G. hederacea is moderately tolerant simultaneously to shade and drought, a characteristic that has been reported for numerous invasive species, while at the same time it shares some characteristics with weedy plants.     

Photosynthetic acclimation in eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings following transfer of shade-grown seedlings to high light

 We studied photosynthetic acclimation of eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings in the first month after sudden exposure of shade-grown seedlings to full sunlight. In a greenhouse experiment, seedlings were grown under full sun or 80% shade, and after 7 months, a sample of the shaded trees was transferred to full sun in the greenhouse. Photosynthetic responses of shaded, transferred, and sun trees were followed over the course of 26 days to track short to medium-term acclimation responses. A partial acclimation of photosynthesis at high light occurred in pre-existing (formed in the previous environment) and new foliage of transferred seedlings. This was associated with non-stomatal limitations to photosynthesis. Pre-existing foliage of transferred plants had a prolonged reduction in the ratio of variable to maximal fluorescence, and a limited capacity to adjust photochemical quenching or photosystem II quantum yield in the light to increasing light intensity compared to sun foliage, and apparently had some difficulty sustaining non-photochemical quenching. Seedling survival was only 58% among transferred seedlings, compared to 80% and 100% in the shade or sun groups, respectively. Photosystem II quantum yield in the light, and photochemical and non-photochemical quenching were similar between newly formed foliage of transferred and sun plants. These findings indicate that eastern hemlock depends strongly on the production of new foliage for photosynthetic adjustments to high light, and that development of photosynthetic competence may be a gradual process that occurs over successive foliar production cycles. Received: 12 May 1998 / Accepted: 27 July 1998     

The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Californian chaparral shrub

The functional roles of the contrasting morphologies of sun and shade shoots of the evergreen shrub Heteromeles arbutifolia were investigated in chaparral and understory habitats by applying a three-dimensional plant architecture simulation model, YPLANT. The simulations were shown to accurately predict the measured frequency distribution of photosynthetic photon flux density (PFD) on both the leaves and a horizontal surface in the open, and gave reasonably good agreement for the more complex light environment in the shade. The sun shoot architecture was orthotropic and characterized by steeply inclined (mean = 71^o) leaves in a spiral phyllotaxy with short internodes. This architecture resulted in relatively low light absorption efficiencies (E _A) for both diffuse and direct PFD, especially during the summer when solar elevation angles were high. Shade shoots were more plagiotropic with longer internodes and a pseudo-distichous phyllotaxis caused by bending of the petioles that positioned the leaves in a nearly horizontal plane (mean = 5^o). This shade-shoot architecture resulted in higher E _A values for both direct and diffuse PFD as compared to those of the sun shoots. Differences in E _A between sun and shade shoots and between summer and winter were related to differences in projection efficiencies as determined by leaf and solar angles, and by differences in self shading resulting from leaf overlap. The leaves exhibited photosynthetic acclimation to the sun and the shade, with the sun leaves having higher photosynthetic capacities per unit area, higher leaf mass per unit area and lower respiration rates per unit area than shade leaves. Despite having 7 times greater available PFD, sun shoots absorbed only 3 times more and had daily carbon gains only double of those of shade shoots. Simulations showed that sun and shade plants performed similarly in the open light environment, but that shade shoots substantially outperformed sun shoots in the shade light environment. The shoot architecture observed in sun plants appears to achieve an efficient compromise between maximizing carbon gain while minimizing the time that the leaf surfaces are exposed to PFDs in excess of those required for light saturation of photosynthesis and therefore potentially photoinhibitory. Received: 8 June 1997 / Accepted: 2 November 1997     

Carboxydismutase Activity in Shade-adapted and Sun-adapted Species of Higher Plants

The activity of the photosynthetic enzyme carboxydismutase (ribulose-l,5-diphosphate carboxylase) was measured in leaf extracts of a number of higher plant species from habitats with greatly contrasting light intensities. Plants occupying sunny habitats and capable of light saturated rates of photosynthesis several times higher than those growing in the deep shade of redwood forests also have a considerably higher carboxydismutase activity. Thus, when expressed on the basis of total chlorophyll or even fresh weight, the enzyme activity is several times greater among the sun than among the shade species. The comparatively low content of soluble protein in the shade plants indicates that their content of enzymes other than carboxydismutase also is low. Nevertheless, the activity of carboxydismutase even on the basis of soluble protein appears to be significantly higher in the sun than in the shade species. It is concluded that low carboxydismutase activity probably is one of the factors that limit the capacity for light saturated photosynthesis in the shade plants.