Growth and water relations of Liquidambar styraciflua L. in an urban park and plaza

Summary Growth and water relations of 10-year-old sweet gum (Liquidambar styraciflua L.) street trees were studied in sites with low and high potential evapotranspiration to determine how these differences are integrated by growth and water relations over time. The trees were located in the parking strip between the curb and sidewalk at a partially vegetated urban park and an urban plaza in Seattle, Washington. Crown size, and seasonal and diurnal stomatal conductance and water potential, as well as diurnal air temperature and humidity, were measured over 2 growing seasons. Yearly trunk growth since transplanting was measured from increment cores. Vapor pressure deficits and air temperatures averaged 18% greater at the plaza, but whole-tree water loss appeared to be much lower than the park trees due to more restricted stomatal conductance and crown size. In addition, yearly diameter increment declined progressively once the plaza trees were established in the existing soil several years after transplanting. Lower water potential in the plaza trees indicated greater internal moisture deficits than the park trees, and tissue analysis revealed lower nutritional status, particularly nitrogen. A manipulative study of water and fertilizer to several additional plaza trees showed an interaction between water and nutrient deficiencies in the coarse and shallow soil that apparently limited growth. Furthermore, soil limitations probably interacted with paved surface conditions over time by reducing nutrient recycling from leaf litter, and generating higher vapor pressure deficits that would contribute to prolonged stomatal closure. Restricted growth and water relations status of the plaza trees represented an equilibrium between chronic high-resource demand above ground and limited below ground.     

Seasonal reversibility of acclimation to irradiance in leaves of common box (Buxus sempervirens L.) in a deciduous forest

Understorey shade plants are seasonally exposed to dramatic changes in light conditions in deciduous forests related with the dynamics of the overstorey leaf phenology. These transitions are commonly followed by changes in herb plant communities, but shade-tolerant evergreen species must be able to adapt to changing light conditions. In this work we checked the photoprotective responses of evergreen species to acclimate to the shady summer environment and reversibly de-acclimate to a more illuminated environment after leaf fall on deciduous overstoreys. For that purpose we have followed the process of light acclimation in leaves of common box (Buxus sempervirens) during the winter to spring transition, which decrease irradiance in the understorey, and conversely during the transition from summer to autumn. Four parameters indicative of the structure and degree of acclimation of the photosynthetic apparatus have been studied: chlorophyll a/b ratio which is supposed to be inversely proportional to the antenna size, α/β-carotene which increases in shade acclimated leaves and the pools of α-tocopherol and xanthophyll cycle pigments (VAZ) which are two of the main photoprotection mechanisms in plants. Among these parameters, chlorophyll a/b ratio and VAZ pool responded finely to changes in irradiance indicating that modifications in the light harvesting size and photoprotective capacity contribute to the continuous acclimation and de-acclimation of long-lived evergreen leaves.     

Is the Lower Shade Tolerance of Scots Pine,Relative to Pedunculate Oak,Related to the Composition of Photosynthetic Pigments?

Foliage of Scots pine (Pinus sylvestris L.) and pedunculate oak (Quercus robur L.) was collected in a mixed pine/oak forest at canopy positions differing in radiation environment. In both species, chlorophyll (Chl) a/b ratios were higher in foliage of canopy positions exposed to higher irradiance as compared to more shaded crown layers. Throughout the growing season, pine needles exhibited significantly lower Chl a/b ratios than oak leaves acclimated to a similar photon availability. Hence, pine needles showed shade-type pigment characteristics relative to foliage of oak. At a given radiation environment, pine needles tended to contain more neoxanthin and lutein per unit of Chl than oak leaves. The differences in pigment composition between foliage of pine and oak can be explained by a higher ratio of outer antennae Chl to core complex Chl in needles of P. sylvestris which enhances the efficiency of photon capture under limiting irradiance. The shade-type pigment composition of pine relative to oak foliage could have been due to a reduced mesophyll internal photon exposure of chloroplasts in needles of Scots pine, resulting from their xeromorphic anatomy. Hence, the higher drought tolerance of pine needles could be achieved at the expense of shade tolerance.     

Relative enhancement of photosynthesis and growth at elevated CO2 is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survivorship of dipterocarp seedlings in the deeply shaded understorey of South‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO₂ upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO₂. Equal total daily photon flux density (PFD) (～7·7 mol m^?2 d^?1) was supplied as either uniform irradiance (～170 µmol m^?2 s^?1) or shade/fleck sequences (～30 µmol m^?2 s^?1/～525 µmol m^?2 s^?1). Photosynthesis and growth were enhanced by elevated CO₂ treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO₂ but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO₂ were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post‐irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO₂ under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.     

Costs and benefits of photosynthetic light acclimation by tree seedlings in response to gap formation

Some shade leaves increase their photosynthetic capacity (P _max) when exposed to a higher irradiance. The increase in P _max is associated with an increase in chloroplast size or number. To accommodate those chloroplasts, plants need to make thick leaves in advance. We studied the cost and benefit of photosynthetic acclimation in mature leaves of a tree species, Kalopanax pictus Nakai, in a cool-temperate deciduous forest. Costs were evaluated as the additional investment in biomass required to make thick leaves, while the benefit was evaluated as an increase in photosynthetic carbon gain. We created gaps by felling canopy trees and examined the photosynthetic responses of mature leaves of the understorey seedlings. In the shade, leaves of K. pictus had vacant spaces that were not filled by chloroplasts in the mesophyll cells facing the intercellular space. When those leaves were exposed to higher irradiance after gap formation, the area of the mesophyll surface covered by chloroplasts increased by 17% and P _max by 27%. This increase in P _max led to an 11% increase in daily carbon gain, which was greater than the amount of biomass additionally invested to construct thicker leaves. We conclude that the capacity of a plant to acclimate to light (photosynthetic acclimation) would contribute to rapid growth in response to gap formation.     

High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survival of dipterocarp seedlings in the understorey of south‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. Field measurements demonstrated that Shorea leprosula seedlings in a rain forest understorey received a high proportion of daily photon flux density at temperatures supra‐optimal for photosynthesis (72% at ≥30 °C, 14% at ≥35 °C). To investigate the effect of high temperatures on photosynthesis during sunflecks, gas exchange and chlorophyll fluorescence measurements were made on seedlings grown in controlled environment conditions either, under uniform, saturating irradiance (approximately 539 µmol m^?2 s^?1) or, shade/fleck sequences (approximately 30 µmol m^?2 s^?1/approximately 525 µmol m^?2 s^?1) at two temperatures, 28 or 38 °C. The rate of light‐saturated photosynthesis, under uniform irradiance, was inhibited by 40% at 38 °C compared with 28 °C. However, during the shade/fleck sequence, photosynthesis was inhibited by 59% at 38 °C compared with 28 °C. The greater inhibition of photosynthesis during the shade/fleck sequence, when compared with uniform irradiance, was driven by the lower efficiency of dynamic photosynthesis combined with lower steady‐state rates of photosynthesis. These results suggest that, contrary to current dogma, sunfleck activity may not always result in significant carbon gain. This has important consequences for seedling regeneration processes in tropical forests as well as for leaves in other canopy positions where sunflecks make an important contribution to total photon flux density.     

INFLUENCE OF NEIGHBORS ON TREE FORM: EFFECTS OF LATERAL SHADE AND PREVENTION OF SWAY ON THE ALLOMETRY OF LIQUIDAMBAR STYRACIFLUA (SWEET GUM)

Liquidambar styraciflua (sweet gum) saplings were open grown (control), guy-wired to prevent wind-induced sway (constrained), or guy-wired within open-topped shade cloth cylinders to simulate shading by neighboring trees (shaded). Mechanical stability was assessed after 2 years of growth by field measurement of whole-tree flexibility and with the critical buckling height predictions of five allometrical models. Data from this experiment are supplemented with measurements of sweet gum trees growing in a 9-year-old pine plantation; individuals crowded but not overtopped by pines are compared with similar-aged trees growing in openings within the plantation. Changes in aboveground architecture were stimulated by both prevention of sway and lateral shade but the effect of the latter was most pronounced; shaded saplings approached or exceeded their calculated critical height and buckled under their own weight when the shade cloth cylinders were removed. Rates of aboveground biomass acquisition, total leaf area, wood density, and wood elastic moduli did not differ among the three treatments; changes in stem allometry accounted for most of the observed differences in tree stability. Sweet gum trees growing in a pine plantation responded similarly to neighbors; crowded trees had larger height: diameter ratios and more closely approached their calculated critical heights than did trees growing in the open. Sweet gum saplings growing in dense stands of similar-sized plants may surmount their neighbors by allocating relatively more wood to height growth than to girth increment.     

Photosynthesis and isoprene emission from trees along an urban–rural gradient in Texas

Isoprene emission is an important mechanism for improving the thermotolerance of plant photosystems as temperatures increase. In this study, we measured photosynthesis and isoprene emission in trees along an urban–rural gradient that serves as a proxy for climate change, to understand daily and seasonal responses to changes in temperature and other environmental variables. Leaf‐level gas exchange and basal isoprene emission of post oak (Quercus stellata) and sweet gum (Liquidambar styraciflua) were recorded at regular intervals over an entire growing season at urban, suburban, and rural sites in eastern Texas. In addition, the temperature and atmospheric carbon dioxide concentration experienced by leaves were experimentally manipulated in spring, early summer, and late summer. We found that trees experienced lower stomatal conductance and photosynthesis and higher isoprene emission, at the urban and suburban sites compared to the rural site. Path analysis indicated a daily positive effect of isoprene emission on photosynthesis, but unexpectedly, higher isoprene emission from urban trees was not associated with improved photosynthesis as temperatures increased during the growing season. Furthermore, urban trees experienced relatively higher isoprene emission at high CO₂ concentrations, while isoprene emission was suppressed at the other sites. These results suggest that isoprene emission may be less beneficial in urban, and potentially future, environmental conditions, particularly if higher temperatures override the suppressive effects of high CO₂ on isoprene emission. These are important considerations for modeling future biosphere–atmosphere interactions and for understanding tree physiological responses to climate change.     

Stem growth and canopy dynamics in a world-wide range of Fagus forests

Abstract. Forests dominated by Fagus (beech) occur widely in the Northern Hemisphere. Tree species dominant together with beech vary in tolerance of understorey conditions. They are deciduous broad-leaved, evergreen broad-leaved or evergreen coniferous. The frequency and intensity of events that reduce the forest canopy cover are important determinants of the ratio of beech to other species in the canopy. For trees in the understorey and the canopy, stem diameter growth rate is determined by light regime and growing space which in turn are determined by canopy cover. We evaluated increase in stem diameter growth rate as an indicator of sudden reductions in canopy cover and canopy dynamics. We used tree-ring chronologies and calculated an index of growth rate increase (GI) to compare the canopy dynamics of 11 natural beech forests. Per site, the annual average value of GI poorly reflected the effects of dry or cool summers, and it clearly reflected events like tornados and hurricanes that removed substantial canopy cover. Among groups of sites average values of GI were significantly different. In the sites with a lower level of average GI, the establishment of the more shade tolerant tree and shrub species in the understorey was favoured, and subcanopy layers became more dense. On the other hand, higher levels of average GI allowed for more light demanding tree species to reach the canopy.     

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     

Photosynthetic acclimation and water-use efficiency of three species of understory herbaceous bamboo (Gramineae) in Panama

Summary To assess the role of photosynthetic acclimation in the response of tropical understory herbs to treefall light gaps, photosynthetic response curves were determined for three species of herbaceous bamboo growing in treatments of sun and shade at Barro Calorado Island, Panama. Increased maximum photosynthetic capacity did not always accompany higher ramet production in the sun treatment. Pharus latifolius reproduced abundantly in both treatments, and produced more ramets and developed higher maximum photosynthetic capacity under higher irradiance. Streptochaeta spicata also produced a high percentage of reproductive ramets in both treatments and produced more ramets in the sun, did not show any significant differences in photosynthetic parameters between treatments. Streptochaeta sodiroana did not change maximum photosynthetic capacity in the sun, and had higher photosynthetic efficiency and lower mortality in the shade. Stable carbon isotope composition of leaves indicated that all three species developed higher water-use efficiency under higher irradiance. Photosynthetic flexibility may contribute to the ability of P. latifolius to reproduce in treefall gaps, whereas S. spicata and S. sodiroana may maintain the ability to fix carbon efficiently in low irradiance even when growing or persisting in gaps.     

Interactions between irradiance,nitrogen nutrition,and water stress in the sun-shade responses of Solanum dulcamara

When grown with adequate water and nitrogen (12 mM NO ₃ ^- ) four clones of Solanum dulcamara from sun or shade habitats in Europe showed similar potential for acclimation of photosynthesis to irradiance level during growth. When grown with limiting nitrogen (0.6 mM NO ₃ ^- ) all clones showed a low potential for acclimation of photosynthesis to irradiance during growth. If limiting nitrogen was accompanied by water stress at high irradiance, the initial slope of the irradiance response curve, and the irradiance saturated rate of photosynthesis were depressed, especially in a clone from a shaded habitat. These interactions are discussed in terms of earlier reports on the sunshade responses and sun-shade ecotypic differentiation in this species.     

光照对两种热带雨林树种幼苗光合能力、热耗散和抗氧化系统的影响

 研究了生长于不同光照条件下（100%、25%和8%光强）热带雨林冠层树种绒毛番龙眼（Pometia tomentosa）和中层树种滇南风吹楠（Horsfieldia tetratepala）幼苗的光合能力、热耗散、活性氧和保护性酶的活性。结果表明,绒毛番龙眼的最大光合速率随着生长光强的增加而提高,而滇南风吹楠在全光条件下的最大光合速率反比25%光照条件下的低。全光条件下两个树种光系统II的最大光化学效率（Fv/Fm）都显著降低,表明发生了长期光抑制。当把生长于遮荫条件下的幼苗移到全光下,从凌晨到中午随着光强的增加光抑制加剧,日落时生长于8%光照条件下的绒毛番龙眼及生长于8%和25%光照条件下的滇南风吹楠的光抑制不能完全恢复。非光化学猝灭对光强的响应曲线表明,随着生长光强的增加滇南风吹楠的热耗散能力增强,而生长在全光和25%光照条件下的绒毛番龙眼的热耗散能力都比滇南风吹楠的弱。两个树种叶片中O-[]·2、H2O2含量、SOD和CAT活性均随着生长光强的增加而提高;在同一光照条件下,绒毛番龙眼叶片中O-[]·2、H2O2含量、SOD和CAT活性显著高于滇南风吹楠。上述结果表明,在光抑制条件下,冠层树种绒毛番龙眼较大程度通过提高保护性酶的活性来保护光合机构免受损伤,而中层树种滇南风吹楠却较大程度通过增强非光化学猝灭来耗散过量光能;滇南风吹楠对强光的适应性差。     

Effects of below- and aboveground competition from the vines Lonicera japonica and Parthenocissus quinquefolia on the growth of the tree host Liquidambar styraciflua

Detrimental effects of vines on tree growth in successional environments have been frequently reported. Little is known, however, about the relative importance of below and aboveground competition from vines on tree growth. The objective of this study was to quantify and compare the growth responses of Liquidambar styraciflua saplings to below and/or aboveground competition with the exotic evergreen vine, Lonicera japonica (Japanese honeysuckle), and the native deciduous vine, Parthenocissus quinquefolia (Virginia creeper). Soil trenching and/or vine-trellising were used to control the type of vine competition experienced by trees. Comparisons among untrenched treatments tested for effects of belowground competition. Comparisons among trenched treatments tested for effects of aboveground competition. After two growing seasons, Lonicera japonica had a greater effect on the growth of L. styraciflua than did P. quinquefolia. This effect was largely due to root competition, as canopy competition only had a negative effect on tree growth when it occurred in combination with root competition. Leaf expansion was consistently and similarly affected by all treatments which involved belowground competition.     

Species-specific acclimation to strong shade modifies susceptibility of conifers to photoinhibition

Photoinhibitory processes in the photosynthetic apparatus of the seedlings of Abies alba (Mill.), Picea abies (Karst.), and Pinus mugo (Turra) growing under strong shade (5 % of full solar irradiance) or full irradiance conditions were investigated in winter and spring using chlorophyll a fluorescence techniques. The extent of photoinhibition in needles as indicated by a decrease in maximum quantum yield of PS II photochemistry (F_v/F_m) depended on species, air temperature and acclimation to the light environment. Unexpectedly, shade-tolerant Abies alba was less affected by low-temperature photoinhibition compared to the other species. F_v/F_m recovered with increasing air temperature. During winter, the seedlings of Picea abies growing in shade showed higher F_v/F_m than those from full light. Non-photochemical quenching of fluorescence (NPQ) measured at the same levels of actinic light was higher in needles acclimated to full light except for Abies alba in February. Photosynthetic performance in term of ETR (apparent electron transfer rate) was also higher in full light-acclimated needles. In April, at ambient temperature, recovery of PS II efficiency from the stress induced by illumination with saturating light was faster in the needles of Picea abies than in those of Abies alba. The shade-acclimated needles of Abies alba and Picea abies showed greater down-regulation of PS II induced by high light stress.     

Early growth and nutritional response to resource competition in cocoa-shade intercropped systems

Intercropping is often promoted for effective mutualism between species, thus compensating for external inputs. However, for optimal farm design resulting in superior production and nutrition, an accurate assessment of plant inter- and intra-specific competition is required. In predominant shade tree-cocoa (Theobroma cacao) systems, inconclusive evidence remains on species interactions, limitations to resource availability and subsequent growth and nutritional response, particularly in early growth. We examined cocoa biomass and foliar nutrition as well as nutrient supply through rates of decomposition and N mineralization after 1-year growth. Our approach employed fertilization and mixed planting treatments in an additive design of cocoa in monoculture (control), under artificial shade, and intercropped under two separate shade species (Terminalia superba and Newbouldia laevis). Intercropping had no effect on cocoa biomass production in comparison to monoculture cocoa. However, artificial shading stimulated foliage and root production both with and without fertilization, suggesting strong effects of light regulation on growth in the absence of belowground competition. Nutritionally, intercropping suppressed K uptake in cocoa foliage as K concentration was reduced by 20–25%, signifying dilution of this nutrient, presumably due to interspecific competition for mobile elements. Foliar N content under N. laevis was raised, where N concentration kept up with growth under this intercropped species. Intercropping also delayed decomposition rates, suggesting slower but sustained release of available nutrients into the topsoil. Cocoa under artificial shade, both with and without fertilization, exhibited the greatest nutrient responses as compared to unfertilized monoculture cocoa, where P uptake was stimulated most (175 and 112%), followed by K (69 and 71%), and then N (54 and 42%). Intercropping with shade trees failed to increase cocoa biomass, however, nutrient uptake was sustained for N and P, suggesting low interspecific competition. When fertilizers are undesirable or unavailable, intercropping of appropriately selected shade trees will not competitively suppress early growth of cocoa but will improve light regulation and nutritional status of cocoa saplings.     

Interplant variation in creosotebush foliage characteristics and canopy arthropods

Summary We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.     

Root production is determined by radiation flux in a temperate grassland community

Accurate knowledge of the response of root turnover to a changing climate is needed to predict growth and produce carbon cycle models. A soil warming system and shading were used to vary soil temperature and received radiation independently in a temperate grassland dominated by Holcus lanatus L. Minirhizotrons allowed root growth and turnover to be examined non‐destructively. In two short‐term (8 week) experiments, root responses to temperature were seasonally distinct. Root number increased when heating was applied during spring, but root death increased during autumnal heating. An experiment lasting 12 months demonstrated that any positive response to temperature was short‐lived and that over a full growing season, soil warming led to a reduction in root number and mass due to increased root death during autumn and winter. Root respiration was also insensitive to soil temperature over much of the year. In contrast, root growth was strongly affected by incident radiation. Root biomass, length, birth rate, number and turnover were all reduced by shading. Photosynthesis in H. lanatus exhibited some acclimation to shading, but assimilation rates at growth irradiance were still lower in shaded plants. The negative effects of shading and soil warming on roots were additive. Comparison of root data with environmental measurements demonstrated a number of positive relationships with photosynthetically active radiation, but not with soil temperature. This was true both across the entire data set and within a shade treatment. These results demonstrate that root growth is unlikely to be directly affected by increased soil temperatures as a result of global warming, at least in temperate areas, and that predictions of net primary productivity should not be based on a positive root growth response to temperature.     

The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand

Variation in light demand is a major factor in determining the growth and survival of trees in a forest. There is strong relation between the light‐demand and the effect of growth irradiance on leaf morphology and photosynthesis in three Acer species: A. rufinerve (light‐demanding), A. mono (intermediate) and A. palmatum (shade‐tolerant). The increase in mesophyll thickness and surface area of chloroplasts facing the intercellular airspaces (S_c) with growth irradiance was highest in A. rufinerve. Although the increase in light‐saturated photosynthesis (A_max) was similar among the species, the increase in water use efficiency (WUE) was much higher in A. rufinerve than that in the other species, indicating that the response to water limitation plays an important role in leaf photosynthetic acclimation to high light in A. rufinerve. The low CO₂ partial pressure at the carboxylation site (C_c) in A. rufinerve (130 µmol mol^?1) at high irradiance was caused by low stomatal and internal conductance to CO₂ diffusion, which minimized the increase in A_max in A. rufinerve despite its high Rubisco content. Under shade conditions, interspecific differences in leaf features were relatively small. Thus, difference in light demand related to leaf acclimation to high light rather than that to low light in the Acer species.     

Limitation of coffee leaf photosynthesis by stomatal conductance and light availability under different shade levels

In agroforestry systems, the effect of shade trees on coffee net photosynthesis (A _n) has been the object of debates among coffee scientists. In this study, we undertook over 600 coffee A _n “spot” measurements under four different artificial shade levels (100, 72, 45 and 19% of full solar irradiance) and analyzed limitations to A _n by low light availability (photon flux density, PFD) and stomatal conductance (g _s). These gas exchange measurements were carried out during two consecutive coffee growing seasons in a commercial plantation in the Orosi valley of Costa Rica. Levels of A _n were related to PFD and g _s in order to calculate envelope functions which were used to establish PFD or g _s limitations to A _n. Under the growing conditions of the present trial, mean leaf A _n remained stable for growth irradiance (GI) as low as 45% of full sun and decreased by ~20% at 19% GI. Limitation to A _n due to g _s was strong in full sun and decreasing with increasing shade levels. On the other hand, limitation due to PFD remained at a similar level for all shade treatments. These different evolutions of limitations of A _n by PFD and g _s in response to shade explain the absence of a decrease in coffee leaf A _n with a shade level up to 55%. Consequently, these results confirm that Arabica coffee is a shade-adapted plant with leaves that can maintain a high photosynthetic performance under low light availability.