Susceptibility of green leaves and green flower petals of CAM orchid <Emphasis Type="Italic">Dendrobium</Emphasis> cv. Burana Jade to high irradiance under natural tropical conditions

Photosynthetic rates of green leaves (GL) and green flower petals (GFP) of the CAM plant Dendrobium cv. Burana Jade and their sensitivities to different growth irradiances were studied in shade-grown plants over a period of 4 weeks. Maximal photosynthetic O₂ evolution rates and CAM acidities [dawn/dusk fluctuations in titratable acidity] were higher in leaves exposed to intermediate sunlight [a maximal photosynthetic photon flux density (PPFD) of 500–600 μmol m⁻² s⁻¹] than in leaves grown under full sunlight (a maximal PPFD of 1 000–1 200 μmol m⁻² s⁻¹) and shade (a maximal PPFD of 200–250 μmol m⁻² s⁻¹). However, these two parameters of GFP were highest in plants grown under the shade and lowest in full sun-grown plants. Both GL and GFP of plants exposed to full sunlight had lower predawn F_v/F_m [dark adapted ratio of variable to maximal fluorescence (the maximal photosystem 2 yield without actinic irradiation)] than those of shade-grown plants. When exposed to intermediate sunlight, however, there were no significant changes in predawn F_v/F_m in GL whereas a significant decrease in predawn F_v/F_m was found in GFP of the same plant. GFP exposed to full sunlight exhibited a greater decrease in predawn F_v/F_m compared to those exposed to intermediate sunlight. The patterns of changes in total chlorophyll (Chl) content of GL and GFP were similar to those of F_v/F_m. Although midday F_v/F_m fluctuated with prevailing irradiance, changes of midday F_v/F_m after exposure to different growth irradiances were similar to those of predawn F_v/F_m in both GL and GFP. The decreases in predawn and midday F_v/F_m were much more pronounced in GFP than in GL under full sunlight, indicating greater sensitivity in GFP to high irradiance (HI). In the laboratory, electron transport rate and photochemical and non-photochemical quenching of Chl fluorescence were also determined under different irradiances. All results indicated that GFP are more susceptible to HI than GL. Although the GFP of Dendrobium cv. Burana Jade require a lower amount of radiant energy for photosynthesis and this plant is usually grown in the shade, is not necessarily a shade plant.     

Photosynthetic response of two tropical liana species grown under different irradiances

We investigated the characteristics of gaseous exchanges and chlorophyll a fluorescence under different irradiances in two liana species Canavalia parviflora Benth. (Fabaceae) and Gouania virgata Reissk (Rhamnaceae), both of a semi-deciduous tropical forest of Southeast Brazil. We used cultivated plants growing under irradiances of 100, 40, 10, and 1.5 % of the photosynthetic photon flux density (PPFD). Higher net photosynthetic rates (P _N) were observed during early morning under full sunlight. After this, reduced P _N values were recorded due to pronounced stomatal closure. In Canavalia, the gas exchange responses diminished concomitant with reduced irradiance. Gouania exhibited a narrower range of response, with high P _N values even at 10 % PPFD. Marked reduction of the effective photochemical yield (ΔF/F_m’) near midday was observed, followed by increases in the non-photochemical quenching for both species under full sunlight. Despite the common occurrence of these species in open areas of the forest, both were able to maintain relatively high P _N in shaded environments. We suggest that lianas present an intermediate physiological behaviour between shade and non-shade tolerant species.     

Light-Induced Adaptive Responses under Greenhouse and Controlled Conditions in the Fern Pteris cretica var. ouvrardii

The photosynthetic capabilities of the fern Pteris cretica var. ouvrardii were analysed by means of the light response curves of CO₂ exchange. In control growth conditions (greenhouse, low-light: 20–32 W m^?2); photosynthesis was shown to be saturated for low irradiance (20–25 W m^?2); the saturating photosynthetic rate, very low as compared to higher plants, was due to an extremely high intracellular resistance. When irradiance during the photosynthesis measurement was higher than 60–80 W m^?2, a constant decline of net CO₂ exchange as a function of time was observed. When irradiance during growth was enhanced, whether in greenhouse (20–250 W m^?2) or controlled (62 W m^?2) conditions, the first fronds that had developed in the new condition from the crosier stage exhibited decreased net maximal photosynthesis and a decreased efficiency in low light, but saturating irradiance was unmodified. However, the fronds whose entire differentiation (from meristem) occurred under these moderate irradiances (plants defoliated of all fronds and crosiers at the time of transfer), possessed more efficient photosynthetic characteristics than control plants. Pteris is able to grow under extreme shade conditions (4–8 W m^?2); light saturating photosynthesis and efficiency are higher under extreme shade than under control conditions. These adaptive characteristics indicate that Pteris is a well-adapted shade species.     

Diurnal changes in chlorophyll fluorescence and light utilization in Colocasia esculenta leaves grown in marshy waterlogged area

Diurnal cycle of chlorophyll fluorescence parameters was done in Colocasia esculenta L. (swamp taro) grown in marshy land under sun or under shade. The sun leaves maintained higher electron transport rate (ETR) and steady state to initial fluorescence ratio (F_s/F₀) than shade leaves. In spite of lower ETR, higher photochemical quenching (PQ), and effective quantum yield of photosystem 2 (Φ_PS2) was evident in shade plants compared to plants exposed to higher irradiance. ETR increased linearly with increase in irradiance more under low irradiance (r ² = 0.84) compared to higher irradiance (r ² = 0.62). The maximum quantum yield of PS 2 (F_v/F_m) did not differ much in sun and shade leaves with the exception of midday when excess of light energy absorbed by plants under sun was thermally dissipated. Hence swamp taro plants adopted different strategies to utilize radiation under different irradiances. At higher irradiance, there was faster decline in proportion of open PS 2 centers (PQ) and excess light energy was dissipated through non-photochemical quenching (NPQ). Under shade, absorbed energy was effectively utilized resulting in higher Φ_PS2.     

An in situ study of photosynthetic oxygen exchange and electron transport rate in the marine macroalga Ulva lactuca (Chlorophyta)

Direct comparisons between photosynthetic O₂ evolution rate and electron transport rate (ETR) were made in situ over 24 h using the benthic macroalga Ulva lactuca (Chlorophyta), growing and measured at a depth of 1.8 m, where the midday irradiance rose to 400–600 μmol photons m⁻² s⁻¹. O₂ exchange was measured with a 5-chamber data-logging apparatus and ETR with a submersible pulse amplitude modulated (PAM) fluorometer (Diving-PAM). Steady-state quantum yield ((F_m′−F_t)/F_m′) decreased from 0.7 during the morning to 0.45 at midday, followed by some recovery in the late afternoon. At low to medium irradiances (0–300 μmol photons m⁻² s⁻¹), there was a significant correlation between O₂ evolution and ETR, but at higher irradiances, ETR continued to increase steadily, while O₂ evolution tended towards an asymptote. However at high irradiance levels (600–1200 μmol photons m⁻² s⁻¹) ETR was significantly lowered. Two methods of measuring ETR, based on either diel ambient light levels and fluorescence yields or rapid light curves, gave similar results at low to moderate irradiance levels. Nutrient enrichment (increases in [NO₃ ⁻], [NH₄ ⁺] and [HPO₄ ^2-] of 5- to 15-fold over ambient concentrations) resulted in an increase, within hours, in photosynthetic rates measured by both ETR and O₂ evolution techniques. At low irradiances, approximately 6.5 to 8.2 electrons passed through PS II during the evolution of one molecule of O₂, i.e., up to twice the theoretical minimum number of four. However, in nutrient-enriched treatments this ratio dropped to 5.1. The results indicate that PAM fluorescence can be used as a good indication of the photosynthetic rate only at low to medium irradiances. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of irradiance on growth,photosynthesis, and water use efficiency of seedlings of the chaparral shrub,Ceanothus megacarpus

Summary The effects of irradiance during growth on biomass allocation, growth rates, leaf chlorophyll and protein contents, and on gas exchange responses to irradiance and CO₂ partial pressures of the evergreen, sclerophyllous, chaparral shrub, Ceanothus megacarpus were determined. Plants were grown at 4 irradiances for the growth experiments, 8, 17, 25, 41 nE cm^-2 sec^-1, and at 2 irradiances, 9 and 50 nE cm^-2 sec^-1, for the other comparisons.At higher irradiances root/shoot ratios were somewhat greater and specific leaf weights were much greater, while leaf area ratios were much lower and leaf weight ratios were slightly lower than at lower irradiances. Relative growth rates increased with increasing irradiance up to 25 nE cm^-2 sec^-1 and then leveled off, while unit leaf area rates increased steeply and unit leaf weight rates increased more gradually up to the highest growth irradiance.Leaves grown at 9 nE cm^-2 sec^-1 had less total chlorophyll per unit leaf area and more per unit leaf weight than those grown at 50 nE cm^-2 sec^-1. In a reverse of what is commonly found, low irradiance grown leaves had significantly higher chlorophyll a/b than high irradiance grown leaves. High irradiance grown leaves had much more total soluble protein per unit leaf area and per unit dry weight, and they had much higher soluble protein/chlorophyll than low irradiance grown leaves.High irradiance grown leaves had higher rates of respiration in very dim light, required higher irradiances for photosynthetic saturation and had higher irradiance saturated rates of photosynthesis than low irradiance grown leaves. CO₂ compensation irradiances for leaves of both treatments were very low, <5 nE cm^-2 sec^-1. Leaves grown under low and those grown under high irradiances reached 95% of their saturated photosynthetic rates at 65 and 85 nE cm^-2 sec^-1, respectively. Irradiance saturated rates of photosynthesis were high compared to other chaparral shrubs, 1.3 for low and 1.9 nmol CO₂ cm^-2 sec^-1 for high irradiance grown leaves. A very unusual finding was that leaf conductances to H₂O were significantly lower in the high irradiance grown leaves than in the low irradiance grown leaves. This, plus the differences in photosynthetic rates, resulted in higher water use efficiencies by the high irradiance grown leaves. High irradiance grown leaves had higher rates of photosynthesis at any particular intercellular CO₂ partial pressure and also responded more steeply to increasing CO₂ partial pressure than did low irradiance grown leaves. Leaves from both treatments showed reduced photosynthetic capability after being subjected to low CO₂ partial pressures (100 bars) under high irradiances. This treatment was more detrimental to leaves grown under low irradiances.The ecological implications of these findings are discussed in terms of chaparral shrub community structure. We suggest that light availability may be an important determinant of chaparral community structure through its effects on water use efficiencies rather than on net carbon gain.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.     

Effects of shading on calcareous benthic periphyton in a short-hydroperiod oligotrophic wetland (Everglades, FL, USA)

The effects of shade on benthic calcareous periphyton were tested in a short-hydroperiod oligotrophic subtropical wetland (freshwater Everglades). The experiment was a split-plot design set in three sites with similar environmental characteristics. At each site, eight randomly selected 1-m² areas were isolated individually in a shade house, which did not spectrally change the incident irradiance but reduced it quantitatively by 0, 30, 50, 60, 70, 80, 90 and 98%. Periphyton mat was sampled monthly under each shade house for a 5 month period while the wetland was flooded. Periphyton was analyzed for thickness, DW, AFDW, chlorophyll a (chl a) and incubated in light and dark BOD bottles at five different irradiances to assess its photosynthesis–irradiance (PI) curve and respiration. The PI curves parameters P _max, I _k and eventually the photoinhibition slope (β) were determined following non-linear regression analyses. Taxonomic composition and total algal biovolume were determined at the end of the experiment. The periphyton composition did not change with shade but the PI curves were significantly affected by it. I _k increased linearly with increasing percent irradiance transmittance (%IT = 1−%shade). P _max could be fitted with a PI curve equation as it increased with %IT and leveled off after 10%IT. For each shade level, the PI curve was used to integrate daily photosynthesis for a day of average irradiance. The daily photosynthesis followed a PI curve equation with the same characteristics as P _max vs. %IT. Thus, periphyton exhibited a high irradiance plasticity under 0–80% shade but could not keep up the same photosynthetic level at higher shade, causing a decrease in daily GPP at 98% shade levels. The plasticity was linked to an increase in the chl a content per cell in the 60–80% shade, while this increase was not observed at lower shade likely because it was too demanding energetically. Thus, chl a is not a good metric for periphyton biomass assessment across variously shaded habitats. It is also hypothesized that irradiance plasticity is linked to photosynthetic coupling between differently comprised algal layers arranged vertically within periphyton mats that have different PI curves.     

Photosynthetic characteristics of ornamental passion flowers grown under different light intensities

Responses of leaf gas exchange, fluorescence emission, chlorophyll concentration, and morpho-anatomical features to changes in photosynthetic photon flux density (PPFD) were studied in three wild ornamental species of Passiflora L. to select sun and shade species for landscaping projects. Artificial shade was obtained with different shading nylon nets, under field conditions, which allowed the reduction of 25, 50, and 75% of global radiation, along with a control treatment under full sunlight. For Passiflora morifolia the highest mean values of light-saturated net photosynthetic rate (P _Nmax) and light compensation point (LCP) were observed at 50 and 25% shade, respectively, while the highest values of dark respiration rate (R _D) and apparent quantum yield (α) were observed at 75% shade. For Passiflora suberosa litoralis the highest value of P _max was observed at full sunlight. The highest mean values for P _max, R _D, and LCP for Passiflora palmeri var. sublanceolata were obtained at 25% shade. The highest values of net photosynthetic rate (P _N) for P. morifolia, P. palmeri var. sublanceolata, and P. suberosa litoralis were 21.09, 16.15, and 12.36 μmol(CO₂) m⁻² s⁻¹, observed at 50 and 75% shade and full sunlight, respectively. The values of the minimal chlorophyll fluorescence (F₀) were significantly different in P. suberosa litoralis and P. palmeri var. sublanceolata, increasing with the increase of the irradiance. In contrast, the values of maximum photochemical efficiency of PSII (F_v/F_m) were significantly different only in P. suberosa litoralis, being higher at 75%, progressively reducing with the increase of PPFD levels. The total concentration of chlorophyll (Chl) was higher in shaded plants than in the ones cultivated in full sunlight. On the other hand, the values of Chl a/b ratio were reduced in shaded plants. A significant effect of shade levels on leaf area (LA) and specific leaf area (SLA) was found for the three species, whose highest mean values were observed at 75% shade. The thickness of foliar tissues was significantly higher for the three species at full sunlight and 25% shade. These results suggested that P. morifolia and P. palmeri var. sublanceolata appeared to be adapted to moderate shade conditions. P. suberosa litoralis presented higher plasticity to greater variation of the irradiance levels, while the photoinhibition was one of the limiting factors for this species at full sunlight.     

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.     

HEAVY METAL‐INDUCED INHIBITION OF PHOTOSYNTHESIS: TARGETS OF IN VIVO HEAVY METAL CHLOROPHYLL FORMATION1

The targets of heavy metal (here Cu^{2 +} and Zn^{2 +} ) attack on the photosynthetic apparatus of algae belonging to different phyla were investigated. Experiments with the green alga Scenedesmus quadricauda confirmed previous findings that according to the irradiance level two different phenomena occur, which were further characterized by specific changes in several photosynthetic parameters. The reaction occurring under low irradiance (shade reaction) is characterized by heavy metal substitution of Mg^{2 +} in chl molecules bound predominantly in the light harvesting complex II of Chlorophyta (LHC II). Under high irradiance (sun reaction) the LHC II chls are inaccessible to substitution and the damage occurs in the PSII reaction center instead. Algae with antenna proteins other than the LHC II did not show the two types of heavy metal attack at different irradiances. In red algae (Antithamnion plumula), low Cu^{2 +} concentrations induced the sun reaction even at very low irradiance. In brown algae (Ectocarpus siliculosus) the shade reaction occurred even in saturating irradiance. These results also indicate that despite some similarity in their features, the primary step of the sun reaction and photoinhibition is different.     

THE SHORT‐TERM EFFECT OF IRRADIANCE ON THE PHOTOSYNTHETIC PROPERTIES OF ANTARCTIC FAST‐ICE MICROALGAL COMMUNITIES1

Although sea‐ice represents a harsh physicochemical environment with steep gradients in temperature, light, and salinity, diverse microbial communities are present within the ice matrix. We describe here the photosynthetic responses of sea‐ice microalgae to varying irradiances. Rapid light curves (RLCs) were generated using pulse amplitude fluorometry and used to derive photosynthetic yield (Φ_PSII), photosynthetic efficiency (α), and the irradiance (E_k) at which relative electron transport rate (rETR) saturates. Surface brine algae from near the surface and bottom‐ice algae were exposed to a range of irradiances from 7 to 262 μmol photons · m^?2 · s^?1. In surface brine algae, Φ_PSII and α remained constant at all irradiances, and rETR_max peaked at 151 μmol photons · m^?2 · s^?1, indicating these algae are well acclimated to the irradiances to which they are normally exposed. In contrast, Φ_PSII, α, and rETR_max in bottom‐ice algae reduced when exposed to irradiances >26 μmol photons · m^?2 · s^?1, indicating a high degree of shade acclimation. In addition, the previous light history had no significant effect on the photosynthetic capacity of bottom‐ice algae whether cells were gradually exposed to target irradiances over a 12 h period or were exposed immediately (light shocked). These findings indicate that bottom‐ice algae are photoinhibited in a dose‐dependent manner, while surface brine algae tolerate higher irradiances. Our study shows that sea‐ice algae are able to adjust to changes in irradiance rapidly, and this ability to acclimate may facilitate survival and subsequent long‐term acclimation to the postmelt light regime of the Southern Ocean.     

Light acclimation, CO2 response and long-term capacity of underwater photosynthesis in three terrestrial plant species

To characterize underwater photosynthetic performance in some terrestrial plants, we determined (i) underwater light acclimation (ii) underwater photosynthetic response to dissolved CO₂, and (iii) underwater photosynthetic capacity during prolonged submergence in three species that differ in submergence tolerance: Phalaris arundinacea, Rumex crispus (both submergence-tolerant) and Arrhenatherum elatius (submergence-intolerant). None of the species had adjusted to low irradiance after 1 week of submergence. Under non-submerged (control) conditions, only R. crispus displayed shade acclimation. Submergence increased the apparent quantum yield in this species, presumably because of the enhanced CO₂ affinity of the elongated leaves. In control plants of the grass species P. arundinacea and A. elatius, CO₂ affinities were higher than for R. crispus. The underwater photosynthetic capacity of R. crispus increased during 1 month of submergence. In P. arundinacea photosynthesis remained constant during 1 month of submergence at normal irradiance; at low irradiance a reduction in photosynthetic capacity was observed after 2 weeks, although there was no tissue degeneration. In contrast, underwater photosynthesis of the submergence-intolerant species A. elatius collapsed rapidly under both irradiances, and this was accompanied by leaf decay. To describe photosynthesis versus irradiance curves, four models were evaluated. The hyperbolic tangent produced the best goodness-of-fit, whereas the rectangular hyperbola (Michaelis-Menten model) gave relatively poor results.     

Photosynthetically versatile thin shade leaves: A paradox of irradiance-response curves

Thick sun leaves have a larger construction cost per unit leaf area than thin shade leaves. To re-evaluate the adaptive roles of sun and shade leaves, we compared the photosynthetic benefits relative to the construction cost of the leaves. We drew photosynthetically active radiation (PAR)-response curves using the leaf-mass-based photosynthetic rate to reflect the cost. The dark respiration rates of the sun and shade leaves of mulberry (Morus bombycis Koidzumi) seedlings did not differ significantly. At irradiances below 250 μmol m⁻² s⁻¹, the shade leaves tended to have a significantly larger net photosynthetic rate (P _N) than the sun leaves. At irradiances above 250 μmol m⁻² s⁻¹, the P _N did not differ significantly. The curves indicate that plants with thin shade leaves have a larger daily CO₂ assimilation rate per construction cost than those with thick sun leaves, even in an open habitat. These results are consistently explained by a simple model of PAR extinction in a leaf. We must target factors other than the effective assimilation when we consider the adaptive roles of thick sun leaves.     

Effects of nitrogen supply on photosynthetic and anatomical changes in current-year needles of <Emphasis Type="Italic">Pinus koraiensis</Emphasis> seedlings grown under two irradiances

We investigated the responses of photon-saturated photosynthesis rate (P _sat) and its simultaneous acclimation of anatomy and nitrogen use patterns of current needles of Korean pine (Pinus koraiensis) seedlings grown under factorial combinations of two nitrogen levels and irradiances. Although N supply resulted in a significant increase of N content in needles under both irradiances, the increase of P _sat tended to be suppressed only in shade (S). The significant increase of P _sat in full sunlight (O) was associated with the increase of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and chlorophyll (Chl) contents. In contrast, small increase of Chl content and no increase of RuBPCO content were found in S (90 % cut of full irradiance), which would result in a small increase of P _sat. This result suggests that extra N is stocked in needles under shade for the growth in next season. With N supply, a significant decrease of specific leaf area (SLA) was detected only in O. This decrease of SLA was due to the increase of density of needle. Furthermore, the increase of needle density was not due to the increased number and size of mesophyll cells, but the increased density of each mesophyll cell. Therefore, although SLA changed in O, the change did not involve anatomical adaptation to use increased N effectively, at least observable by light microscopy. Hence, even though the SLA would change, N deposition will improve the photosynthetic capacity of Korean pine seedlings, not through the development of needle anatomy but through improvement of the allocation of N in both irradiances.     

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.     

PHOTOSYNTHESIS IN AN EXTREME SHADE ENVIRONMENT: BENTHIC MICROBIAL MATS FROM LAKE HOARE, A PERMANENTLY ICE-COVERED ANTARCTIC LAKE

We investigated the composition of benthic microbial mats in permanently ice-covered Lake Hoare, Antarctica, and their irradiance vs. photosynthetic oxygen exchange relationships. Mats could be subdivided into three distinct depth zones: a seasonally ice-free “moat” zone and two under-ice zones. The upper under-ice zone extended from below the 3.5 m thick ice to approximately 13 m and the lower from below 13 m to 22 m. Moat mats were acclimated to the high irradiance they experienced during summer. They contained photoprotective pigments, predominantly those characteristic of cyanobacteria, and had high compensation and saturating irradiances (E_c and E_k) of 75 and 130 μmol photons·m⁻²·s⁻¹, respectively. The moat mats used light inefficiently. The upper under-ice community contained both cyanobacteria and diatoms. Within this zone, biomass (as pigments) increased with increasing depth, reaching a maximum at 10 m. Phycoerythrin was abundant in this zone, with shade acclimation and efficiency of utilization of incident light increasing with depth to a maximum of 0.06 mol C fixed·mol⁻¹ incident photons under light-limiting conditions. Precipitation of inorganic carbon as calcite was associated with this community, representing up to 50% of the carbon sequestered into the sediment. The lower under-ice zone was characterized by a decline in pigment concentrations with depth and an increasing prevalence of diatoms. Photosynthesis in this community was highly shade acclimated and efficient, with E_c and E_k below 0.5 μmol·m⁻²·s⁻¹ and 2 μmol·m⁻²·s⁻¹, respectively, and maximum yields of 0.04 mol C fixed·mol⁻¹ incident quanta. Carbon uptake in situ by both under-ice and moat mats was estimated at up to 100 and 140 mg·m⁻²·day⁻¹, based on the photosynthesis–irradiance curves, incident irradiance, and light attenuation by ice and the water column.     

The photochemical reflectance index: an optical indicator of photosynthetic radiation use efficiency across species, functional types, and nutrient levels

The photochemical reflectance index (PRI), derived from narrow-band reflectance at 531 and 570 nm, was explored as an indicator of photosynthetic radiation use efficiency for 20 species representing three functional types: annual, deciduous perennial, and evergreen perennial. Across species, top-canopy leaves in full sun at midday exhibited a strong correlation between PRI and ΔF/Fm′, a fluorescence-based index of photosystem II (PSII) photochemical efficiency. PRI was also significantly correlated with both net CO₂ uptake and radiation use efficiency measured by gas exchange. When species were examined by functional type, evergreens exhibited significantly reduced midday photosynthetic rates relative to annual and deciduous species. This midday reduction was associated with reduced radiation use efficiency, detectable as reduced net CO₂ uptake, PRI, and ΔF/Fm′ values, and increased levels of the photoprotective xanthophyll cycle pigment zeaxanthin. For each functional type, nutrient deficiency led to reductions in both PRI and ΔF/Fm′ relative to fertilized controls. Laboratory experiments exposing leaves to diurnal courses of radiation and simulated midday stomatal closure demonstrated that PRI changed rapidly with both irradiance and leaf physiological state. In these studies, PRI was closely correlated with both ΔF/Fm' and radiation use efficiency determined from gas exchange at all but the lowest light levels. Examination of the difference spectra upon exposure to increasing light levels revealed that the 531 nm Δ reflectance signal was composed of two spectral components. At low irradiance, this signal was dominated by a 545-nm component, which was not closely related to radiation use efficiency. At progressively higher light levels above 100 μmol m⁻² s⁻¹, the 531-nm signal was increasingly dominated by a 526-nm component, which was correlated with light use efficiency and with the conversion of the xanthophyll pigment violaxanthin to antheraxanthin and zeaxanthin. Further consideration of the two components composing the 531-nm signal could lead to an index of photosynthetic function applicable over a wide range of illumination. The results of this study support the use of PRI as an interspecific index of photosynthetic radiation use efficiency for leaves and canopies in full sun, but not across wide ranges in illumination from deep shade to full sun. The discovery of a consistent relationship between PRI and photosynthetic radiation use efficiency for top-canopy leaves across species, functional types, and nutrient treatments suggests that relative photosynthetic rates could be derived with the “view from above” provided by remote reflectance measurements if issues of canopy and stand structure can be resolved. Received: 6 January 1997 / Accepted: 14 July 1997     

Midday photoinhibition of two newly developed super-rice hybrids

Super-rice hybrids are two-line hybrid rice cultivars with 15 to 20 % higher yields than the raditional three-line hybrid rice cultivars. Response of photosynthetic functions to midday photoinhibition was compared between seedlings of the traditional hybrid rice (Oryza sativa L.) Shanyou63 and two super-rice hybrids, Hua-an3 and Liangyoupeijiu. Under strong midday sunlight, in comparison with Shanyou63, the two super-rice hybrids were less photoinhibited, as indicated by the lower loss of the net photosynthetic rate (P _N), the quantum yield of photosystem 2 (Φ_PS2), and the maximum and effective quantum yield of PS2 photochemistry (F_v/F_m and F_v′/F_m′). They also had a much higher transpiration rate. Hence the super-rice hybrids could protect themselves against midday photoinhibition at the cost of water. The photoprotective de-epoxidized xanthophyll cycle components, antheraxanthin (A) and zeaxanthin (Z), were accumulated more in Hua-an3 and Liangyoupeijiu than in Shanyou63, but the size of xanthophyll cycle pool of the seedlings was not affected by midday photoinhibition. Compared to Shanyou63, the super-rice hybrids were better photoprotected under natural high irradiance stress and the accumulation of Z and A, not the size of the xanthophyll pool protected the rice hybrids against photoinhibition.     

Comparisons of photosynthesis and photoinhibition in the CAM vine Hoya australis and several C3 vines growing on the coast of eastern Australia

Abstract. The CAM vine Hoya australis and three C₃ vines, Smilax australis, Ipomoea pes-caprae and Kennedta rubicunda, were studied at a site on the coast of northeastern New South Wales, Australia. The level of CAM activity (nocturnal acid accumulation) was comparable in H. australis growing in full sunlight and in deep shade. Acclimation to shade by H. australis was indicated by thinner leaves, increased chlorophyll content, decreased chlorophyll a/b ratios, lower dark respiration rates, and lower light compensation points. When growing in full sunlight H. australis exhibited reductions in photochemical efficiency, as indicated by reduced quantum yields and F_v/F_m fluorescence from PS II as well as low rates of photosynthesis at high light. Sun leaves of H. australis experienced a massive quenching of fluorescence from PS II during normal exposure to midday irradiance which was rapidly reversible under low irradiance conditions in the late afternoon. This quenching indicated a reduction in photochemical efficiency, part of which could be accounted for by an increase in non-radiative energy dissipation, while part of it was due to one or more processes not yet identified. Changes in PS II fluorescence from shade H. australis exposed to full sunlight suggest a decrease in the rate constant for photochemistry indicative of damage to the reaction centre, as well as an increase in non-radiative energy dissipation. The C₃ vine S. australis was also shade tolerant, but exhibited little evidence of photoinhibition when growing in full sunlight. Ipomoea pescaprae and K. rubicunda, both of which were apparently shade intolerant (being found only in full sunlight), possessed high quantum yields and much higher photosynthetic capacities than either H. australis or S. australis. From this study, and several others, it appears that plants possessing CAM experience photoinhibition to a greater degree than do C₃ species in full sunlight under natural conditions, which is probably exacerbated by some degree of CAM-idling.