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     

Tradeoffs Between Irradiance Capture and Avoidance in Semi-arid Environments Assessed with a Crown Architecture Model

Plants in arid environments cope with stress from excessiveirradiance by physiological photoprotection of the photosyntheticapparatus and by structurally reducing the leaf area exposedto the sun (structural photoprotection). We assessed the ecologicalrelevance of structural photoprotection in two plants of contrastingarchitecture co-occurring in a semi-arid environment, usingthe three-dimensional canopy model YPLANT. We compared the roleof crown geometry in avoiding excessive radiation, analysedthe costs of structural photoprotection in terms of reductionof potential carbon gain, and compared these costs with thosedue to seasonal constraints of photosynthesis and tissue ageing.The results of the model simulations indicated that canopy architectureofStipa tenacissima(a tussock grass) andRetama sphaerocarpa(aleafless leguminous shrub) minimized the risk of overheatingand photo-oxidative destruction of the photosynthetic apparatuswith steeply oriented foliage and moderate self-shading. Butthis structural photoprotection imposed an increased cost interms of potential carbon gain. Diurnal and seasonal patternsof light interception by the crown of these plants translatedinto a simulated potential carbon gain only half that of anequivalent, horizontal photosynthetic surface. This reductionin potential carbon gain, due to irradiance avoidance, was similarto that imposed by water shortage.S. tenacissima,which ceasesphotosynthetic activity during periods of drought, exhibitedmore structural avoidance of irradiance thanR. sphaerocarpa,whichremains active throughout the year. This illustrates the influenceof the capacity of plants to utilize light for carbon fixationon the trade-offs between irradiance capture and avoidance.Structural avoidance of excessive radiation efficiently preventsthe risk of damage by intense irradiance, has no special maintenancecosts, and is biomechanically cheaper than enhanced light harvestingby a horizontal canopy, which points to structural photoprotectionas a very effective strategy to cope with high irradiance stressin poor and adverse habitats.Copyright 1999 Annals of BotanyCompany Canopy model, functional architecture, irradiance avoidance, light harvesting, photoinhibition, photoprotection, photosynthesis model, plant architecture,Retama sphaerocarpa,semi-arid environment,Stipa tenacissima,structural photoprotection.     

The geometry of light interception by shoots of Heteromeles arbutifolia: morphological and physiological consequences for individual leaves

The influence of leaf orientation and position within shoots on individual leaf light environments, carbon gain, and susceptibility to photoinhibition was studied in the California chaparral shrub Heteromeles arbutifolia with measurements of gas exchange and chlorophyll fluorescence, and by application of a three-dimensional canopy architecture model. Simulations of light absorption and photosynthesis revealed a complex pattern of leaf light environments and resulting leaf carbon gain within the shoots. Upper, south-facing leaves were potentially the most productive because they intercepted greater daily photon flux density (PFD) than leaves of any other orientation. North-facing leaves intercepted less PFD but of this, more was received on the abaxial surface because of the steep leaf angles. Leaves differed in their response to abaxial versus adaxial illumination depending on their orientation. While most had lower photosynthetic rates when illuminated on their abaxial as compared to adaxial surface, the photosynthetic rates of north-facing leaves were independent of the surface of illumination. Because of the increasing self-shading, there were strong decreases in absorbed PFD and daily carbon gain in the basipetal direction. Leaf nitrogen per unit mass also decreased in the basipetal direction but on a per unit area basis was nearly constant along the shoot. The decrease in leaf N per unit mass was accounted for by an increase in leaf mass per unit area (LMA) rather than by movement of N from older to younger leaves during shoot growth. The increased LMA of older lower leaves may have contributed directly to their lower photosynthetic capacities by increasing the limitations to diffusion of CO₂ within the leaf to the sites of carboxylation. There was no evidence for sun/shade acclimation along the shoot. Upper leaves and especially south-facing upper leaves had a potential risk for photoinhibition as demonstrated by the high PFDs received and the diurnal decreases in the fluorescence ratio F _v/F _m. Predawn F _v/F _m ratios remained high (>0.8) indicating that when in their normal orientations leaves sustained no photoinhibition. Reorientation of the leaves to horizontal induced a strong sustained decrease in F _v/F _m and CO₂ exchange that slowly recovered over the next 10–15?days. If leaves were also inverted so that the abaxial surface received the increased PFDs, then the reduction in F _v/F _m and CO₂ assimilation was much greater with no evidence for recovery. The heterogeneity of responses was due to a combination of differences between leaves of different orientation, differences between responses on their abaxial versus adaxial surfaces, and differences along the shoot due to leaf age and self-shading effects.     

Ontogeny, understorey light interception and simulated carbon gain of juvenile rainforest evergreens differing in shade tolerance

Background and Aims

A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.

Methods

The biomass distribution of juveniles (17–740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.

Results

An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR_d = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.

Conclusions

The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.     

Sexes show contrasting patterns of leaf and crown carbon gain in a dioecious rainforest shrub

The sexes of dioecious species may differ in a range of vegetative and reproductive traits as well as in physiological traits. In Siparuna grandiflora, a Neotropical dioecious shrub, we examined differences in leaf-level photosynthesis of different classes of leaf age and, using simulation models, explored whether differences in leaf-level carbon gain led to sex differences in whole-plant daily carbon gain. Male plants had higher photosynthetic capacity at the leaf level. As leaves of both sexes aged their photosynthetic capacity and specific leaf area declined as expected. Simulations of daily carbon gain using the architecturally explicit model Y-Plant and a non-architectural model incorporating a wide range of realistic light environments revealed that the difference in leaf-level photosynthetic capacity did not translate into greater crown-level carbon gain for males. Rather, differences in patterns of allocation to leaf area allow females to achieve higher crown-level carbon gain. The results demonstrate that sex differences at the leaf level do not necessarily predict patterns at the whole-plant level.     

A relationship between carbon dioxide, photosynthetic efficiency and shade tolerance

Net photosynthesis and transpiration of seedlings from shade tolerant, moderately tolerant and intolerant tree species were measured in ambient carbon dioxide (CO₂) concentrations ranging from 312 to 734 ppm. The species used, Fagus grandifolia Ehrh. (tolerant), Quercus alba L., Q. rubra L., Liriodendron tulipifera L. (moderately tolerant), Liquidambar styraciflua L. and Pinus taeda L. (intolerant), are found co-occurring in the mixed pine-hardwood forests of the Piedmont region of the southeastern United States. When seedlings were grown in shaded conditions, photosynthetic CO₂ efficiency was significantly different in all species with the highest efficiency in the most shade tolerant species, Fagus grandifolia , and progressively lower efficiencies in moderately tolerant and intolerant species. Photosynthetic CO₂ efficiency was defined as the rate of increase in net photosynthesis with increase in ambient CO₂ concentration. When plants which had grown in a high light environment were tested, the moderately tolerant and intolerant deciduous species had the highest photosynthetic CO₂ efficiencies but this capacity was reduced when these species grew in low light. The lowest CO₂ efficiency and apparent quantum yield occurred in Pinus taeda in all cases. Water use efficiency was higher for all species in enriched CO₂ environments but transpiration rate and leaf conductance were not affected by CO₂ concentration. High photosynthetic CO₂ efficiency may be advantageous for maintaining a positive carbon balance in the low light environment under a forest canopy.     

Leaf display and photosynthesis of tree seedlings in a cool-temperate deciduous broadleaf forest understorey

To examine a possible convergence in leaf photosynthetic characteristics and leaf display responses to light environment in seedlings of three canopy and two shrub tree species in understorey of cool-temperate deciduous broadleaf forest, relationships between light environment, leaf orientation and leaf light-photosynthetic response were measured. Light capture of the seedlings (17-24 individuals with 2-12 leaves for each species) was assessed with a three dimensional geometric modeling program Y-plant. Leaf photosynthetic characteristics of the five species were found to have acclimated to the understorey light environment, i.e., low light compensation point and high apparent quantum yield. In addition, light-saturated photosynthetic rates were higher in seedlings inhabiting microsites with higher light availability. Efficiencies of light capture and carbon gain of the leaf display were evaluated by simulating the directionalities of light capture and daily photosynthesis for each seedling using hemispherical canopy photography. The results showed that most of the seedlings orientated their leaves in a way to increase the daily photosynthesis during the direct light periods (sunflecks) rather than maximize daily photosynthesis by diffuse light. Simulations also showed that daily photosynthesis would increase only 10% of that on actual leaf display when the leaves orientated to maximize the diffuse light interception. Simulations in which leaf orientations were varied showed that when the leaf display fully maximized direct light interception, the time that leaves were exposed to excessive photon flux density of >800 mumol photons m(-2) s(-1) were doubled. The understorey seedlings studied responded to the given light environments in a way to maximize the efficiency of acquisition and use of light during their short (approximately 3 month) seasonal growth period.     

Needle age dependence of photosynthesis along a light gradient within an <Emphasis Type="Italic">Abies alba</Emphasis> crown

Age and shade effects on needle structure and photosynthesis were determined within a lower part of Abies alba crown along a horizontal increasing gradient of branches self-shading. It was hypothesized that a decrease in net CO₂ assimilation rate with increasing needles’ age would be related to: (1) structural age-related changes in needles, (2) reduction of stomatal conductance, (3) nitrogen translocation from old to young needles, and (4) decrease in efficiency of photochemical processes. Leaf mass-to-area ratio increased non-linearly with needle age. In a needle cross section, distance between the vascular bundles decreased, and height of palisade parenchyma cells increased with age. The structural changes observed in our study might lead to an increase in internal resistance to CO₂ with greater needle age. Total needle nitrogen concentration linearly decreased with age due to dilution and/or translocation to younger needles. When expressed per needle area, nitrogen content was reduced in 6-year-old needles compared with younger ones. Net CO₂ assimilation rate per needle area decayed and was accompanied by a decrease in transpiration and water and photosynthetic nitrogen use efficiency. Old needles maintained high photochemical efficiency which compensated to some extent for light deficit in their micro-light environments. Our results have suggested that there is a mechanism controlling the relation between efficiency of light and dark photosynthetic processes along the needle age and shade gradient in A. alba crown.     

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.     

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.     

Photosynthetic light response in three carnivorous plant species: <Emphasis Type="Italic">Drosera rotundifolia,D. capensis and Sarracenia leucophylla</Emphasis>

Photosynthetic properties of carnivorous plants have not been well characterized and the extent to which photosynthesis contributes to carbon gain in most carnivorous plants is also largely unknown. We investigated the photosynthetic light response in three carnivorous plant species, Drosera rotundifolia L. (sundew; circumpolar and native to northern British Columbia, Canada), Sarracenia leucophylla Rafin. (‘pitcher-plant’; S.E. United States), and D. capensis L. (sundew; Cape Peninsula, South Africa), using portable gas-exchange systems to explore the capacity for photosynthetic carbon gain in carnivorous plant species. Maximal photosynthetic rates (1.32–2.22 μmol m⁻² s⁻¹ on a leaf area basis) and saturating light intensities (100 to 200 μmol PAR m⁻² s⁻¹) were both low in all species and comparable to shade plants. Field or greenhouse-grown D. rotundifolia had the highest rates of photosynthesis among the three species examined. Dark respiration, ranging from −1.44 (S. leucophylla) to −3.32 (D. rotundifolia) μmol m⁻² s⁻¹ was high in comparison to photosynthesis in the species examined. Across greenhouse-grown plants, photosynthetic light compensation points scaled with light-saturated photosynthetic rates. An analysis of gas-exchange and growth data for greenhouse-grown D. capensis plants suggests that photosynthesis can account for all plant carbon gain in this species.     

Stomatal and photosynthetic responses to variable sunlight

Most plants experience many fluctuations in sunlight from full sun to shade throughout the day. Under these conditions, stomatal and photosynthetic responses vary dramatically among species depending on water status and growth form. Many herbaceous, fast-growing species rapidly reduce stomatal opening during short-term shade periods. Rapid stomatal closure during shade conserves water, but may also reduce CO₂ uptake. Because periods of alternating sun and shade can reduce accumulative water stress that would otherwise severely curtail carbon gain, some herbs are restricted to habitats with intermittent periods of shade. In contrast to herbaceous growth forms, woody species maintain relatively constant stomatal opening during both sun and shade periods. This results in greater CO₂ uptake, but with greater water loss. These two generalized response patterns for woody and herbaceous species to natural variations in sunlight conflict with conventional ideas of water use and carbon gain based on measurements made under constant light.     

A hemiparasite in the forest understorey: photosynthetic performance and carbon balance of Melampyrum pratense

• Melampyrum pratense is an annual root‐hemiparasitic plant growing mostly in forest understorey, an environment with unstable light conditions. While photosynthetic responses of autotrophic plants to variable light conditions are in general well understood, light responses of root hemiparasites have not been investigated.
• We carried out gas exchange measurements (light response and photosynthetic induction curves) to assess the photosynthetic performance of M. pratense in spring and summer. These data and recorded light dynamics data were subsequently used to model carbon balance of the hemiparasite throughout the entire growth season.
• Summer leaves had significantly lower rates of saturated photosynthesis and dark respiration than spring leaves, a pattern expected to reflect the difference between sun‐ and shade‐adapted leaves. However, even the summer leaves of the hemiparasite exhibited a higher rate of light‐saturated photosynthesis than reported in non‐parasitic understorey herbs. This is likely related to its annual life history, rare among other understorey herbs. The carbon balance model considering photosynthetic induction still indicated insufficient autotrophic carbon gain for seed production in the summer months due to limited light availability and substantial carbon loss through dark respiration.
• The results point to potentially high importance of heterotrophic carbon acquisition in M. pratense, which could be of at least comparable importance as in other mixotrophic plants growing in forests – mistletoes and partial mycoheterotrophs. It is remarkable that despite apparent evolutionary pressure towards improved carbon acquisition from the host, M. pratense retains efficient photosynthesis and high transpiration rate, the ecophysiological traits typical of related root hemiparasites in the Orobanchaceae.

     

A functional analysis of the crown architecture of tropical forest<Emphasis Type="Italic"> Psychotria</Emphasis> species: do species vary in light capture efficiency and consequently in carbon gain and growth?

The crown architectures of 11 Psychotria species native to Barro Colorado Island, Panama were reconstructed from field measurements of leaf and branch geometry with the three-dimensional simulation model Y-plant. The objective was to assess the role of species differences in architecture in light capture and carbon gain in their natural understory environment. When species were grouped according to their putative light environment preference, the shade tolerant species were found to have a small but significantly higher efficiency of light capture for both diffuse and direct light as compared to the light demanding species. Within each grouping, however, there were few significant differences in light capture efficiency among species. The lower efficiencies of light demanding species was due to slightly higher self-shading and slightly lower angular efficiencies. Simulations of whole plant assimilation showed that light demanding species had greater daily assimilation in both direct and diffuse light due to the significantly greater light availability in the sites where light demanding species were found, as compared to those where shade tolerant species occurred. Among light demanding species, the above ground relative growth rate measured over a 1-year period by applying allometric equations for mass versus linear dimensions, was positively correlated with diffuse PFD and with mean daily assimilation estimated from Y-plant. For the shade tolerant plants, there was no significant correlation between RGR and mean daily assimilation or with any measure of light availability, probably because they occurred over a much narrower range of light environments. Overall, the results reveal a strong convergence in light capture efficiencies among the Psychotria species at lower values than previously observed in understory plants using similar approaches. Constraints imposed by other crown functions such as hydraulics and biomechanical support may place upper limits on light capture efficiency.Abbreviations E_a Efficiency of light absorption (dimensionless) - E_adir Efficiency of direct light absorption (dimensionless) - E_adif Efficiency of diffuse light absorption (dimensionless) - D_E Display efficiency (dimensionless) - P_E Projection efficiency (dimensionless) - CosI Mean cosine of incidence (dimensionless) - aLAR_e Effective leaf area ratio (m² g^–1) - A_tot Daily assimilation (mmol m^–2 day^–1) - A_dir Daily assimilation in direct PFD (mmol m^–2 day^–1) - A_dif Daily assimilation in diffuse PFD (mmol m^–2 day^–1)     

Stomatal and photosynthetic responses to shade in sorghum, soybean and eastern gamagrass

We studied photosynthetic and stomatal responses of grain sorghum ( Sorghum bicolor [L.] Moench cv. Pioneer 8500), soybean ( Glycine max L. cv. Flyer) and eastern gamagrass ( Tripsacum dactyloides L.) during experimental sun and shade periods simulating summer cloud cover. Leaf gas exchange measurements of field plants showed that short-term (5 min) shading of leaves to 300–400 μmol m⁻² s⁻¹ photosynthetic photon flux density reduced photosynthesis, leaf temperature, stomatal conductance, transpiration and water use efficiency and increased intercellular CO₂ partial pressure. In all species, photosynthetic recovery was delayed when leaves were reilluminated, apparently by stomatal closure. The strongest stomatal response was in soybean. Photosynthetic recovery was studied further with soybeans grown indoors (maximum photosynthetic photon flux density 1 200 μmol m⁻² s⁻¹). Plants grown indoors had responses to shade similar to those of field plants, except for brief nonstomatal limitation immediately after reillumination. These responses indicated the importance of the light environment during leaf development on assimilation responses to variable light, and suggested different limitations on carbon assimilation in different parts of the soybean canopy. Photosynthetic oxygen evolution recovered immediately upon reillumination, indicating that the light reactions did not limit soybean photosynthetic recovery. While shade periods caused stomatal closure and reduced carbon gain and water loss in all species, the consequences for carbon gain/water loss were greatest in soybean. The occurrence of stomatal closure in all three species may arise from their shared phenologies and herbaceous growth forms.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

The study of a determinate growth orchid highlights the role of new leaf production in photosynthetic light acclimation

Plants usually respond to the changes of growth irradiance by a combination of the physiological modifications in their preexisting leaves and the production of new leaves. However, those with a determinate growth habit produce certain number of leaves in a growing season and cannot produce new leaves when light condition changes. We used an epiphytic orchid with only one leaf produced every growing season to examine whether and how determinate growth species adapt to changing environments after their preexisting leaves mature. Leaf photosynthesis and anatomy of Pleione aurita were investigated at full expansion and at 40 days after the fully expanded leaves were transferred from high to low light or from low to high light. Leaves show large physiological and morphological plasticity to light gradients at full expansion and the transferred leaves exhibited multiple physiological modifications, including reallocation of nitrogen between light harvesting and carbon fixation, and enhancement of thermal dissipation in their new environments, to optimize carbon assimilation and avoid photoinhibition. Irrespective of the various changes either to shade or sun, the sole preexisting leaf could not fully acclimate to new light environments due to the mesophyll thickness constraint. This leads to the consequence that only plants exposed to high light throughout the experiment had a positive annual biomass gain. Our results highlighted the importance of new leaf production in the carbon accumulation during photosynthetic light acclimation, and contribute new insights of epiphytes physiological responses to their highly dynamic arboreal habitat.     

Photosynthetic acclimation to variability in the light environment of early and late successional plants

Summary Fourteen plant species from early-, mid-, and late-successional habitats were grown for a period of 25 to 50 days in each of two light environments, i.e. full sunlight and in deep shade. The rate of photosynthesis for newly formed leaves was measured as a function of light intensity for plants from each light environment. Photosynthetic flexibility, measured as the difference in response between sun- and shade-grown plants, was determined for each of 5 parameters including dark respiration, quantum yield, light compensation, half-saturating irradiance for photosynthesis, and the photosynthetic rate at 1,400 E m^-2 s^-1. We found photosynthetic flexibility to be high for early successional annuals, intermediate for midsuccessional species, and low for late successional species.     

Photosynthetic and genetic differentiation in sun and shade populations of Holcus mollis L.

Three Holcus mollis L. populations, one with 2n = 28 chromosomes living in a forest and two with 2n = 35 chromosomes, the first living in a forest, the second in open land, are compared for photosynthesis.Simultaneous measurements of oxygen and carbon dioxide, either in high light, low light, or dark experiments indicate that the 2n = 28 chromosomes population is photosynthetically well adapted to shade, while 2n = 35 chromosomes forest population, is not.The 2n = 35 chromosomes plants growing in the forest does not automatically acquire the photosynthetic character of a shade plant, the genome must show an evolution for this. In our study, only the plants with 2n = 28 chromosomes demonstrated the shade adaptation.     

植株叶片的光合色素构成对遮阴的响应

叶绿素在植株体内负责光能的吸收、传递和转化, 类胡萝卜素则行使光能捕获和光破坏防御两大功能, 它们在光合作用中起着非常重要的作用。该文综述了几大主要光合色素的分布和功能, 以及不同物种的色素含量和构成差异。阳生植物的叶黄素库较大, 但脱环氧化水平不及阴生植物。黄体素与叶黄素库的比值与植物的耐阴性呈正相关关系。由不同的遮阴源造成的遮阴环境, 光强和光质有很大的差异, 总体来说对植物生长的影响, 建筑物遮阴<阔叶林遮阴<针叶林遮阴。光强的改变可诱导类胡萝卜素的两大循环——叶黄素循环和黄体素循环。由光强诱导的叶绿素含量和叶绿素a/b比值的改变与该物种的耐阴性无关。短时间的遮阴不会对植物的生长造成危害, 叶黄素库的大小不仅与每天接受的光量子有关, 更与光量子在一天的分布有关, 因为光照和温度是协同作用的。光合作用或色素构成是蓝光、红光和远红光共同作用的结果, 不是某一种单色光所能替代的。我们总结了影响植物色素构成的内因和外因, 指出植物主要通过调整光反应中心和捕光天线色素蛋白复合体的比例, 以及两个光系统的比值来调整色素含量和构成以适应不同的光照条件, 提出了现存研究中存在的一些问题, 旨在为今后的相关研究提供建议。