Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species

Leaf photosynthetic CO₂ responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO₂ assimilation rates (A_net). However, triose‐phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower A_net in high [CO₂] and low [O₂] than at ambient [O₂]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of A_net at saturating [CO₂] (i.e. A_max) to 2 kPa O₂. Three species showed >20% reductions in A_max in low [O₂], a phenomenon potentially explained by orthophosphate (P_i) savings during photorespiration. These species, with largest photosynthetic capacity and P_i > 2 mmol P m^?2, rely the most on additional P_i made available from photorespiration rather than species growing in P‐impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting A_max and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered.     

Aboveground biomass allocation,leaf growth,and photosynthesis patterns in tundra plant forms in arctic Alaska

Summary Tundra plant growth forms can generally be characterized as consisting predominantly of low-growing perennial grasses and sedges, perennial herbaceous forbs, dwarf deciduous shrubs, and dwarf evergreen shrubs. Gross aboveground carbon allocation, leaf growth, and photosynthesis pattern studies were initiated to develop a quantitative understanding of the functional importance of these particular tundra growth forms. Photosynthetic capacities of 13 species were determined under standardized exposure conditions using a¹⁴CO₂ field system and ranged between 5 and 47 mg CO₂·g dry wt^-1·h^-1. These results, in conjunction with detailed leaf growth determinations, support the generalization that species with an evergreen growth form have lower photosynthetic capacities than species with a perennial graminoid, forb, or deciduous shrub growth form. However, these low photosynthetic capacities in evergreen shrubs are associated with relatively extended leaf longevities. Conversely, deciduous shrub forms exhibited high photosynthetic capacities, but were offset by relatively short leaf longevity periods. The perennial grasses, sedges, and forbs showed patterns intermediate to these. As a result, it appears that among tundra species of different growth form, photosynthetic capacity is inversely related to leaf longevity.     

HIGH PHOTOSYNTHETIC RATES UNDER A COLIMITATION FOR INORGANIC PHOSPHORUS AND CARBON DIOXIDE1

Inorganic phosphorus (P_i) and carbon (here, CO₂) potentially limit the photosynthesis of phytoplankton simultaneously (colimitation). A single P_i limitation generally reduces photosynthesis, but the effect of a colimitation is not known. Therefore, photosynthesis was measured under P_i‐limited conditions and high and low CO₂, and osmo‐mixotrophic (i.e., growth in the presence of glucose) conditions that result in colimiting conditions in some cases. The green alga Chlamydomonas acidophila Negoro was used as a model organism because low P_i and CO₂ concentrations likely influence its photosynthetic rates in its natural environment. Results showed a decreasing maximum photosynthetic rate (P_max) and maximum quantum yield (Φ_II) with increasing P_i limitation. In addition, a P_i limitation enhanced the relative contribution of dark respiration to P_max (R_d:P_max) but did not influence the compensation light intensity. P_max positively correlated with the cellular RUBISCO content. Osmo‐mixotrophic conditions resulted in similar P_max, Φ_II, and RUBISCO content as in high‐CO₂ cultures. The low‐CO₂ cultures were colimited by P_i and CO₂ and had the highest P_max, Φ_II, and RUBISCO content. Colimiting conditions for P_i and CO₂ in C. acidophila resulted in an enhanced mismatch between photosynthesis and growth rates compared to the effect of a single P_i limitation. Primary productivity of colimited phytoplankton could thus be misinterpreted.     

No photosynthetic down-regulation in sweetgum trees (Liquidambar styraciflua L.) after three years of CO2 enrichment at the Duke Forest FACE experiment

Photosynthetic capacity and leaf properties of sun and shade leaves of overstorey sweetgum trees (Liquidambar styraciflua L.) were compared over the first 3 years of growth in ambient or ambient + 200 μL L^?1 CO₂ at the Duke Forest Free Air CO₂ Enrichment (FACE) experiment. We were interested in whether photosynthetic down‐regulation to CO₂ occurred in sweetgum trees growing in a forest ecosystem, whether shade leaves down‐regulated to a greater extent than sun leaves, and if there was a seasonal component to photosynthetic down‐regulation. During June and September of each year, we measured net photosynthesis (A) versus the calculated intercellular CO₂ concentration (C_i) in situ and analysed these response curves using a biochemical model that described the limitations imposed by the amount and activity of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Vc_max) and by the rate of ribulose‐1,5‐bisphosphate (RuBP) regeneration mediated by electron transport (J_max). There was no evidence of photosynthetic down‐regulation to CO₂ in either sun or shade leaves of sweetgum trees over the 3 years of measurements. Elevated CO₂ did not significantly affect Vc_max or J_max. The ratio of Vc_max to J_max was relatively constant, averaging 2·12, and was not affected by CO₂ treatment, position in the canopy, or measurement period. Furthermore, CO₂ enrichment did not affect leaf nitrogen per unit leaf area (N_a), chlorophyll or total non‐structural carbohydrates of sun or shade leaves. We did, however, find a strong relationship between N_a and the modelled components of photosynthetic capacity, Vc_max and J_max. Our data over the first 3 years of this experiment corroborate observations that trees rooted in the ground may not exhibit symptoms of photosynthetic down‐regulation as quickly as tree seedlings growing in pots. There was a strong sustained enhancement of photosynthesis by CO₂ enrichment whereby light‐saturated net photosynthesis of sun leaves was stimulated by 63% and light‐saturated net photosynthesis of shade leaves was stimulated by 48% when averaged over the 3 years. This study suggests that this CO₂ enhancement of photosynthesis will be sustained in the Duke Forest FACE experiment as long as soil N availability keeps pace with photosynthetic and growth processes.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

Water- and nutrient-use efficiency of a deciduous species, Vaccinium myrtillus, and an evergreen species, V. vitis-idaea, in a subalpine dwarf shrub heath in the southern Alps, Italy

Periodic measurements of gas‐exchange rates and determinations of foliar N and P concentrations were used for evaluating instantaneous water‐use efficiency and photosynthetic nutrient‐use efficiency in two co‐existing dwarf shrubs of different growth form (V. myrtillus, deciduous, and V. vitis‐idaea, evergreen) in a subalpine heath in the southern Alps of Italy. Those data were compared with cumulative assessments of water‐use efficiency and photosynthetic nutrient‐use efficiency obtained by measuring leaf carbon isotope discrimination in leaf tissues and by estimating nutrient resorption from senescing leaves. V. myrtillus presented higher dry‐weight based rates of net photosynthesis (A_weight) compared to V. vitis‐idaea. A_weight was positively correlated with foliar‐nutrient status and intercellular‐to‐ambient gradient in CO₂ concentrations. A_weight was, furthermore, negatively correlated with leaf specific mass. Instantaneous photosynthetic nutrient‐use efficiency did not differ between the two species but the percentages of N and P pools resorbed from senescing leaves were somewhat higher in the deciduous species. The evergreen species showed lower P concentrations in senescing leaves which indicated a higher proficiency in resorbing phosphorus compared to the deciduous species. In addition, the evergreen species achieved a higher carbon gain per unit foliar N and P, due to a longer mean residence time of both nutrients. The two species did not differ from each other with respect to both instantaneous and long‐term water‐use efficiency. This was consistent with the climatic pattern, showing no sign of water deficiency through the growing season. Current‐year V. vitis‐idaea leaves had a significantly higher Δ¹³C compared to previous‐year leaves, possibly mirroring a long term acclimation of evergreen leaves, as far as they age, to the habitat conditions in the understory where evergreen species are usually confined within mixed dwarf‐shrub communities.     

Limitation to Carbon Assimilation of Two Perennial Species in Semi-Arid South-East Spain

Diurnal and seasonal changes of net photosynthetic rate (P_n) and the efficiency of photosystem 2 (F_v/F_m) were measured on two perennial species growing on a soil catena in semi-arid south-east Spain. Stipa tenacissima, a tussock grass, grows on shallow soil at the top of the catena and Retama sphaerocarpa, a leguminous shrub, grows in the valley bottom. A linear relationship was found between light saturated photosynthetic rates (P_max) and diffusive leaf conductance (g_l) in both Retama and Stipa indicating that the intercellular CO₂ concentration (c_i) was maintained constant in both species diurnally. Relatively high values of calculated c_i in Retama cladodes suggested that was not the primary limitation to carbon assimilation. F_v/F_m for the two species when well watered was around 0.8. Although Retama cladodes maintained this value throughout the year, F_v/F_m decreased to a minimum of 0.43 in Stipa leaves, at the end of the dry season. Our data suggest that plants in the Rambla Honda can substantially reduce transpiration without reducing photosynthetic rates to the same extent by closing their stomata, because P_n is reduced primarily by high respiration, decreased mesophyll conductance and by photoinhibition or permanent damage of photosystem 2.     

Effects of continuous leaf wetness on photosynthesis: adverse aspects of rainfall

Above-ground parts of Phaseolus vulgaris L. plants were treated with artificial misty rain (‘rain’) in a growth chamber to investigate the effects of leaf wetness on photosynthetic performance. The following results were obtained. (1) Stomata closed completely within 2 min of the onset of continuous ‘rain’ application and gradually opened to half the original aperture by 60 min. The rate of CO₂ exchange measured on such wet leaves changed in parallel with the changes in stomatal aperture and attained 60 to 70% of the control level by 1h. (2) The dependence of the rate of leaf photosynthesis, A, on the intercellular CO₂ concentration, c_i [A(c_i) relationship], examined in thoroughly dried leaves which had been treated with ‘rain’ did not change until after 4 h of treatment. However, leaves treated for 6h showed discernible decreases in A at high c_i (c_i>500μmolmol ^?1). The photosynthetic rate of leaves treated with ‘rain’ for 24 h was reduced at all c_i, and A at the ambient CO₂ concentration of 350μmolmol^?1 was 60 to 70% of that of the control level. The rate of photosynthesis did not recover even after 3 d of treatment of the plants in a dry environment. These results clearly indicate that leaf wetness causes not only instantaneous suppression of photosynthesis but also chronic damage to the photosynthetic apparatus. Potential effects of leaf wetness on photosynthetic performance in nature are also discussed.     

The use of low [CO2] to estimate diffusional and non-diffusional limitations of photosynthetic capacity of salt-stressed olive saplings

In this study it has been shown that increased diffusional resistances caused by salt stress may be fully overcome by exposing attached leaves to very low [CO₂] (～ 50 µmol mol^?1), and, thus a non‐destructive‐in vivo method to correctly estimate photosynthetic capacity in stressed plants is reported. Diffusional (i.e. stomatal conductance, g_s, and mesophyll conductance to CO₂, g_m) and biochemical limitations to photosynthesis (A) were measured in two 1‐year‐old Greek olive cultivars (Chalkidikis and Kerkiras) subjected to salt stress by adding 200 mm NaCl to the irrigation water. Two sets of A–C_i curves were measured. A first set of standard A–C_i curves (i.e. without pre‐conditioning plants at low [CO₂]), were generated for salt‐stressed plants. A second set of A–C_i curves were measured, on both control and salt‐stressed plants, after pre‐conditioning leaves at [CO₂] of ～ 50 µmol mol^?1 for about 1.5 h to force stomatal opening. This forced stomata to be wide open, and g_s increased to similar values in control and salt‐stressed plants of both cultivars. After g_s had approached the maximum value, the A–C_i response was again measured. The analysis of the photosynthetic capacity of the salt‐stressed plants based on the standard A–C_i curves, showed low values of the J_max (maximum rate of electron transport) to V_cmax (RuBP‐saturated rate of Rubisco) ratio (1.06), that would implicate a reduced rate of RuBP regeneration, and, thus, a metabolic impairment. However, the analysis of the A–C_i curves made on pre‐conditioned leaves, showed that the estimates of the photosynthetic capacity parameters were much higher than in the standard A–C_i responses. Moreover, these values were similar in magnitude to the average values reported by Wullschleger (Journal of Experimental Botany 44, 907–920, 1993) in a survey of 109 C₃ species. These findings clearly indicates that: (1) salt stress did affect g_s and g_m but not the biochemical capacity to assimilate CO₂ and therefore, in these conditions, the sum of the diffusional resistances set the limit to photosynthesis rates; (2) there was a linear relationship (r² = 0.68) between g_m and g_s, and, thus, changes of g_m can be as fast as those of g_s; (3) the estimates of photosynthetic capacity based on A–C_i curves made without removing diffusional limitations are artificially low and lead to incorrect interpretations of the actual limitations of photosynthesis; and (4) the analysis of the photosynthetic properties in terms of stomatal and non‐stomatal limitations should be replaced by the analysis of diffusional and non‐diffusional limitations of photosynthesis. Finally, the C₃ photosynthesis model parameterization using in vitro‐measured and in vivo‐measured kinetics parameters was compared. Applying the in vivo‐measured Rubisco kinetics parameters resulted in a better parameterization of the photosynthesis model.     

The “one‐point method” for estimating maximum carboxylation capacity of photosynthesis: A cautionary tale

The maximum carboxylation capacity of Rubisco, V_c,max, is an important photosynthetic parameter that is key to accurate estimation of carbon assimilation. The gold‐standard technique for determining V_c,max is to derive V_c,max from the initial slope of an A–C_i curve (the response of photosynthesis, A, to intercellular CO₂ concentration, C_i). Accurate estimates of V_c,max derived from an alternative and rapid “one‐point” measurement of photosynthesis could greatly accelerate data collection and model parameterization. We evaluated the practical application of the one‐point method in six species measured under standard conditions (saturating irradiance and 400 μmol CO₂ mol^?1) and under conditions that would increase the likelihood for successful estimation of V_c,max: (a) ensuring Rubisco‐limited A by measuring at 300 μmol CO₂ mol^?1 and (b) allowing time for acclimation to saturating irradiance prior to measurement. The one‐point method significantly underestimated V_c,max in four of the six species, providing estimates 21%–32% below fitted values. We identified ribulose‐1,5‐bisphosphate‐limited A, light acclimation, and the use of an assumed respiration rate as factors that limited the effective use of the one‐point method to accurately estimate V_c,max. We conclude that the one‐point method requires a species‐specific understanding of its application, is often unsuccessful, and must be used with caution.     

Light‐induced systemic signalling down‐regulates photosynthetic performance of soybean leaves with different directional effects

• When plants are exposed to a heterogeneous environment, photosynthesis of leaves is not only determined by their local condition, but also by certain signals from other parts of the same plant, termed systemic regulation. Our present study was conducted to investigate the effects of light‐dependent systemic regulation on the photosynthetic performance of soybean (Glycine max L. Merr.) under heterogeneous light conditions.
• Soybean plants were treated with heterogeneous light. Then gas exchange characteristics were measured to evaluate the photosynthetic performance of leaves. Parameters related to photosynthetic pigments, chlorophyll fluorescence, Rubisco and photosynthates were examined to study the mechanisms of light‐dependent systemic regulation on photosynthesis.
• Light‐induced systemic signalling by illuminated leaves reduced the P_n of both upper and lower non‐illuminated leaves on the same soybean plant. The decrease in g_s and increase in C_i in these non‐illuminated leaves indicated restriction of carbon assimilation, which was further verified by the decline in content and activity of Rubisco. However, the activation state of Rubisco decreased only in upper non‐illuminated leaves. Quantum efficiency of PSII (ΦPSII) and ETR also decreased only in upper non‐illuminated leaves. Moreover, the effects of light‐induced systemic signalling on carbohydrate content were also detectable only in upper non‐illuminated leaves.
• Light‐induced systemic signalling by illuminated leaves restricts carbon assimilation and down‐regulates photosynthetic performance of non‐illuminated leaves within a soybean plant. However, effects of such systemic regulation differed when regulated in upward or downward direction.

     

An In Situ Leaf and Branch Warming Experiment in the Amazon

Leaves and branches of mature trees, lianas, and gap species were warmed in an Amazonian forest for 4 mo to observe the effect of warming on photosynthesis, stomatal conductance, and transpiration. Electric resistance heaters increased air temperatures near the leaves by approximately 2°C. Sunlit leaf temperatures increased by 2–3°C on average, but during some periods leaf temperatures increased by >5°C. Maximum photosynthesis (A_max) decreased significantly in the warmed leaves vs. the control leaves over the 13‐wk study period with an average decrease in A_max of 1.4 μmol/m²s (19% decrease from a mean A_max of 7.2 μmol/m²s) when measured at 30°C and there were no signs of acclimation to higher temperatures within existing leaves. The decline in A_max was likely due to irreversible temperature damage caused by very high leaf temperatures and not due to C_i limitation of carboxylation. Warming had a larger negative impact on A_max in canopy level tree species than other tested functional groups such as lianas or gap species. Transpiration did not significantly increase in the warmed leaves compared with the control group. This study indicates that increased temperatures due to global warming could potentially decrease future tropical forest carbon uptake by a significant amount. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Photosynthetic characteristics of dwarf and fringe Rhizophora mangle L. in a Belizean mangrove

Twin Cays (Belize) is a highly oligotrophic mangrove archipelago dominated by Rhizophora mangle L. Ocean‐fringing trees are 3–7 m tall with a leaf area index (LAI) of 2.3, whereas in the interior, dwarf zone, trees are 1.5 m or less, and the LAI is 0.7. P‐fertilization of dwarf trees dramatically increases growth. As a partial explanation of these characteristics, it was hypothesized that differences in stature and growth rates would reflect differences in leaf photosynthetic capacity, as determined by the photochemical and biochemical characteristics at the chloroplast level. Gas exchange and chlorophyll fluorescence were used to compare photosynthesis of dwarf, fringe and fertilized trees. Regardless of zonation or treatment, net CO₂ exchange (A) and photosynthetic electron transport were light saturated at less than 500 µmol photons m^?2 s^?1, and low‐light quantum efficiencies were typical for healthy C₃ plants. On the other hand, light‐saturated A was linearly related to stomatal conductance (g_s), with seasonal, zonal and treatment differences in photosynthesis corresponding linearly to differences in the mean g_s. Overall, photosynthetic capacity appeared to be co‐regulated with stomatal conductance, minimizing the variability of C_i at ambient CO₂ (and hence, C_i/C_a). Based on the results of in vitro assays, regulation of photosynthesis in R. mangle appeared to be accomplished, at least in part, by regulation of Rubisco activity.     

Factors underlying genotypic differences in the induction of photosynthesis in soybean [Glycine max (L.) Merr.]

Crop leaves are subject to continually changing light levels in the field. Photosynthetic efficiency of a crop canopy and productivity will depend significantly on how quickly a leaf can acclimate to a change. One measure of speed of response is the rate of photosynthesis increase toward its steady state on transition from low to high light. This rate was measured for seven genotypes of soybean [Glycine max (L.) Merr.]. After 10 min of illumination, cultivar ‘UA4805’ (UA) had achieved a leaf photosynthetic rate (P_n) of 23.2 μmol · m^?2 · s^?1, close to its steady‐state rate, while the slowest cultivar ‘Tachinagaha’ (Tc) had only reached 13.0 μmol · m^?2 · s^?1 and was still many minutes from obtaining steady state. This difference was further investigated by examining induction at a range of carbon dioxide concentrations. Applying a biochemical model of limitations to photosynthesis to the responses of P_n to intercellular CO₂ concentration (C_i), it was found that the speed of apparent in vivo activation of ribulose‐1:5‐bisphosphate carboxylase/oxygenase (Rubisco) was responsible for this difference. Sequence analysis of the Rubisco activase gene revealed single nucleotide polymorphisms that could relate to this difference. The results show a potential route for selection of cultivars with increased photosynthetic efficiency in fluctuating light.     

SEASONAL PATTERNS OF PHOTOSYNTHESIS AND LIGHT-INDEPENDENT CARBON FIXATION IN MARINE MACROPHYTES1

The contribution of light-independent carbon fixation (LICF) to the overall carbon gain and the seasonal patterns of maximum photosynthesis (P_max and LICF were characterized in a broad taxonomic range of macrophytes from Monterey Bay, California. P_max and LICF rates (nmol C.g filtered seawater^?1.min^?1) varied among species and taxonomic groups examined, and as a function of tissue type in the phaeophyte Laminaria setchellii Silva (Phaeophyceae). On average, P_max values were higher in the Rhodophyta, whereas LICF rates were greater in the Phaeophyceae. LICF rates were generally less than 5% of P_max in the marine macrophytes studied and, as a consequence, cannot fully compensate for respiratory carbon losses, which usually are greater than 10% of P_max. All species studied possessed the highest P_max and LICF rates when irradiance levels were highest and decreased during periods of low incident irradiance. Seasonal patterns of P_max and LICF in most of the macrophytes from the stenothermal environment of Monterey Bay were strongly correlated with photosynthetic photon flux rather than seawater temperature. The concomitant decrease of LICF and P_max rates in all species examined argues against LICF playing a major role in carbon acquisition under light-limiting conditions as suggested previously. Rather, the strong positive correlation of P_max and LICF indicates the direct coupling of photosynthate (e.g. 3-phosphoglyceric acid) generation with production of substrates for LICF reactions. Our results also suggest that LICF might be a useful indicator of photosynthetic metabolism in marine macrophytes.     

Leaf Photosynthesis of the Mangrove Avicennia Germinans as Affected by NaCl

In leaves of the mangrove species Avicennia germinans (L.) L. grown in salinities from 0 to 40 ‰, fluorescence, gas exchange, and δ¹³C analyses were done. Predawn values of F_v/F_m were about 0.75 in all the treatments suggesting that leaves did not suffer chronic photoinhibition. Conversely, midday F_v/F_m values decreased to about 0.55-0.60 which indicated strong down-regulation of photosynthesis in all treatments. Maximum photosynthetic rate (P _max) was 14.58 ± 0.22 μmol m^-2 s^-1 at 0 ‰ it decreased by 21 and 37 % in plants at salinities of 10 and 40 ‰, respectively. Stomatal conductance (g _s) was profoundly responsive in comparison to P _max which resulted in a high water use efficiency. This was further confirmed by δ¹³C values, which increased with salinity. From day 3, after salt was removed from the soil solution, P _max and g _s increased up to 13 and 30 %, respectively. However, the values were still considerably lower than those measured in plants grown without salt addition. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic acclimation in relation to nitrogen allocation in cucumber leaves in response to changes in irradiance

Leaves deep in canopies can suddenly be exposed to increased irradiances following e.g. gap formation in forests or pruning in crops. Studies on the acclimation of photosynthesis to increased irradiance have mainly focused on the changes in photosynthetic capacity (A_max), although actual irradiance often remains below saturating level. We investigated the effect of changes in irradiance on the photosynthesis irradiance response and on nitrogen allocation in fully grown leaves of Cucumis sativus. Leaves that fully developed under low (50 µmol m^?2 s^?1) or moderate (200 µmol m^?2 s^?1) irradiance were subsequently exposed to, respectively, moderate (LM‐leaves) or low (ML‐leaves) irradiance or kept at constant irradiance level (LL‐ and MM‐leaves). Acclimation of photosynthesis occurred within 7 days with final A_max highest in MM‐leaves, lowest in LL‐leaves and intermediate in ML‐ and LM‐leaves, whereas full acclimation of thylakoid processes underlying photosystem II (PSII) efficiency and non‐photochemical quenching occurred in ML‐ and LM‐leaves. Dark respiration correlated with irradiance level, but not with A_max. Light‐limited quantum efficiency was similar in all leaves. The increase in photosynthesis at moderate irradiance in LM‐leaves was primarily driven by nitrogen import, and nitrogen remained allocated in a similar ratio to Rubisco and bioenergetics, while allocation to light harvesting relatively decreased. A contrary response of nitrogen was associated with the decrease in photosynthesis in ML‐leaves. Net assimilation of LM‐leaves under moderate irradiance remained lower than in MM‐leaves, revealing the importance of photosynthetic acclimation during the leaf developmental phase for crop productivity in scenarios with realistic, moderate fluctuations in irradiance that leaves can be exposed to.     

LIGHT ABSORPTION BY PLANTS AND ITS IMPLICATIONS FOR PHOTOSYNTHESIS

The preceding account has attempted to examine the interactions between light absorption and photosynthesis, with reference to both unicellular and multicellular terrestrial and aquatic plants. There are, however, some notable plant groups to which no direct reference has been made, e.g. mosses, liverworts and lichens. Although many have similar optical properties to terrestrial vascular plants (Gates, 1980) and apparently similar photosynthetic responses (see Green & Snelgar, 1982; Kershaw, 1984) they may possess subtle, as yet unknown differences. For instance, the lichen thallus has a high surface reflectance although the transmittance is virtually zero (Gates, 1980; Osborne, unpublished results). It is envisaged, however, that differences in optical properties between species will reflect differences in degree not kind. Although not all variation in photosynthesis is due to differences in light absorption a number of accounts suggest that this is a contributing factor. Variations in leaf absorptance have been found to account for most of the variation in leaf photosynthesis at low J_is (see Ehleringer & Björkman, 1978a; Osborne & Garrett, 1983). There is, however, little direct experimental evidence on light absorption and photosynthesis in either microalgal species or aquatic macrophytes. We also do not know over what range of incident photon flux densities photosynthesis is determined largely by changes in light absorption. Plants growing under natural conditions also experience large diurnal and seasonal fluctuations in J_i, unlike species grown under laboratory conditions. The occurrence of transitory peaks in J_i tends to overshadow the fact that the average J_i is often lower than the J₁ required to saturate photosynthesis, i.e. 1500–2000 μmol m^-2 s^-1, depending on the growth treatment. Using the data of Monteith (1977) and I W m²= 5 μmol m^-2 s^-1, and with photosynthetically active radiation 50% of total solar radiation, the daily mean value for Britain is approximately 450 μmol m^-2 s^-1, with a maximum in June of 1000μmol m^-2 s^-1 and a minimum during the winter of 75 μmol m^-2 s^-1. Such values could be even lower on shaded understory leaves and considerably lower for aquatic species. Based on average values of net photosynthesis for a terrestrial plant leaf, light saturation would only be expected in June while for most of the year the average values would lie largely on the light-limited portion of the photosynthesis light response curve. Although the daily average values in tropical climates may be higher during the winter months, they are remarkably similar throughout the world for the respective summers in the northern and southern hemispheres, because the increased daylength at high latitudes compensates for the lower J_is. The expected lower dark respiration rates during the winter may also partially offset the effects of a lower light level. There is therefore a trade-off between high J_is for a short period of time against a lower J_i for a longer period of time. We might expect different photosynthetic responses to these two very different conditions. Importantly, a low J_i with a long daylength may enable a plant to photosynthesize at or near its maximum photon efficiency for most of the day. Although the response of the plant to fluctuations in J_i is complicated because it is affected by the previous environmental conditions, this may indicate that light absorption has a much greater significance under natural conditions, particularly for perennial species. The bias in many laboratories towards research on terrestrial vascular plants also tends to ignore the fact that a number of multicellular and unicellular aquatic species survive in very low light environments. Furthermore, the direct extrapolation of photosynthetic responses from measurements on single leaves to those of whole plants is clearly erroneous. Although this is obvious, many physiological ecologists have attributed all manner of things to the photosynthetic responses of ‘primary’ leaves. Most researchers have ignored problems associated with composite plant tissues and internal light gradients. Clearly caution is required in interpreting the photosynthesis light-response curve of multicellular tissues based on biochemical features alone. Also, the importance of cell structure on light absorption and photosynthesis has generally been ignored and attributed solely to the effects of structural features on CO₂ diffusion. In doing so the work of two or three generations of plant physiologists has been ignored. Haberlandt (1914) at the turn of the century probably first implicated the role of cell structure in leaf optics, and Heath (1970) stressed that in order to completely understand the role of light in photosynthesis we need to know the flux incident on the chloroplast itself. Even this suggestion may need modification because of the capacity of the internal chloroplast membranes for scattering light. It is worth emphasizing the importance of light gradients within tissues and their role in regulating photosynthesis, particularly at light saturation. Measurements of light gradients are fraught with problems because of experimental difficulties and the majority (few) are based on reflectance and transmittance measurements. Seyfried & Fukshansky (1983) have shown that light incident on the lower surface of a Cucurbita cotyledon produced a larger light gradient than light incident from above, indicating the importance of the spatial arrangement of the tissues with respect to the light source. Also, light incident on the lower surface of leaves of Picea sitchensis was less ‘effective’ in photosynthesis than light from above (Leverenz & Jarvis, 1979). Clearly, two tissues could have the same gross absorptance but different photosynthetic rates because of differences in the internal light environment. Fisher & Fisher (1983) have recently found asymmetries in the light distribution within leaves, which they related to asymmetries in photosynthetic products due to differences in solar elevation. Such modifications in light distribution could be important for a number of solar-tracking species. Changes in light absorption are brought about by a whole gamut of physiological, morphological and behavioural responses which serve to optimize the amount of light absorbed. Perhaps the simplest way of regulating the amount of light absorbed is by restricting growth either to particular times of the year or to conditions when the light climate is favourable. We are still largely ignorant of many details of these modifications. In particular, differences in tissue structure such as the size and number of vacuoles or the effects of organelles on the scattering component of the internal light environment of photosynthetic tissues are not understood. A better understanding of the interaction of light with plants in aquatic systems is also required. It is unfortunate that light-absorptance measurements are not routinely made in photosynthetic studies, and this is quite clearly a neglected area of study. That these measurements are not made is even more surprising, since techniques have been available for over sixty years (Ulbricht, 1920). Absorptance measurements are of particular importance in the photosynthetic adaptation of microalgae, where only a small proportion of the incident photon flux density is absorbed. For multicellular species more detailed information is required on internal light gradients and their variability. Light-absorptance measurements are also important in any study relating kinetic data on CO₂ fixation to in vivo photosynthesis, especially when there are large variations in the morphology and structure of the photosynthetic organ.     

Response of photosynthesis of different plant functional types to environmental changes along Northeast China Transect

Net photosynthesis (P_n), transpiration (E), stomatal conductance (g_s), internal CO₂ concentration (C_i), and water use efficiency (WUE) were examined on 215 species from eight plant functional types (PFTs) along a precipitation gradient in northeast China (the Northeast China Transect, or NECT). Among the eight PFTs, meadow steppe grasses had the highest rates of net photosynthesis and forest grasses the lowest and the following order of P_n was noted: meadow steppe grasses >typical steppe grasses >steppe shrubs >desert grasses >forest trees >forest shrubs >desert shrubs >forest grasses (P<0.05). Transpiration tended to be the highest in the steppe grasses and lowest in forest shrubs. Transpiration also decreased rapidly with the appearance of C₃ desert species at the desert end. The forest tree PFT had lower P_n, E, g_s than the steppe PFTs, whereas WUE values were somewhat greater in the forest tree PFT than the desert shrubs and grasses. Low C_i values along the steppe section (from 400 to 1100 km, east to west) indicated the presence of C₄ species. Of all the PFTs, only shrubs and herbs were noted at all points along the transect. No clear relationship between P_n, E, g_s, WUE of herb and shrub PFTs and annual precipitation was noted – low values were found at both the high and low precipitation ends of the transect. Highest values were noted when precipitation was intermediate. Received: 28 October 1998 / Accepted: 10 May 1999     

Andreaea Hartmanii Thed.—A histological study

Abstract

Photosynthetic responses to light intensity were studied under laboratory conditions in seven bryophyte species from evergreen laurel forest, a threatened habitat, on Terceira island in the Azores. Four mosses (Andoa berthelotiana, Echinodium prolixum, Fissidens serrulatus, Myurium hochstetteri) and three liverworts (Bazzania azorica, Frullania tamarisci, Lepidozia cupressina) were selected to encompass a range of potential responses to variations in the forest light environment. Carbon dioxide exchange measurements were made, using an infra-red gas-analyser, at photosynthetic photon flux densities (PPFD) of 0-900 µmol m^-2 s^-1 and a mean temperature of 21°C in fully hydrated shoots. Most species achieved light saturation of photosynthesis below 30 µmol m^-2 s^-1, the lowest value being for A. berthelotiana (20 µmol m^-2 s^-1) and the highest for M. hochstetteri (68 µmol m^-2 s^-1). The liverwort F. tamarisci had the highest maximum photosynthetic rate (P_max, 23 µmol CO₂ g^-1 h^-1) whereas P_max was lowest in the mosses E. prolixum and M. hochstetteri (10 µmol CO₂ g^-1 h^-1). Dark respiration rate, a critical factor in toleration of shade by forest floor plants, was highest in the species with the highest values for P_max. Compensation point was extremely low (7 µmol photons m^-2 s^-1) in Fissidens serrulatus, a species found in the deep shade of forest ravines and caves, and highest in M. hochstetteri a moss restricted to better illuminated habitats within and outside the forest. No photoinhibition was detected during the relatively short exposures to high irradiances. Comparison of these responses with data on the forest light environment indicates that, despite the possession of considerable shade adaptations, during winter in the evergreen laurel forest, low light levels may often limit photosynthetic rates of the bryophytes.