Relations between electron transport rates determined by pulse amplitude modulated chlorophyll fluorescence and oxygen evolution in macroalgae under different light conditions

The relationship between O₂-based gross photosynthesis (GP) and in vivo chlorophyll fluorescence of Photosystem II-based electron transport rate (ETR) as well as the relationship between effective quantum yield of fluorescence (Φ_PSII) and quantum yield of oxygen evolution (Φ_{O_2}) were examined in the green algae Ulva rotundata and Ulva olivascens and the red alga Porphyra leucosticta collected from the field and incubated for 3 days at 100 μmol m⁻² s⁻¹ in nutrient enriched seawater. Maximal GP was twice as high in Ulva species than that measured in P. leucosticta. In all species ETR was saturated at much higher irradiance than GP. The initial slope of ETR versus absorbed irradiance was higher than that of GP versus absorbed irradiance. Only under absorbed irradiances below saturation or at values of GP <2 μmol O₂ m⁻² s⁻¹ a linear relationship was observed. In the linear phase, calculated O₂ evolved /ETR molar ratios were closed to the theoretical value of 0.25 in Ulva species. In P. leucosticta, the estimated GP was associated to the estimated ETR only at high irradiances. ETR was determined under white light, red light emitting by diodes and solar radiation. In Ulva species the maximal ETR was reached under red light and solar radiation whereas in P. leucosticta the maximal ETR was reached under white light and minimal under red light. These results are in agreement with the known action spectra for photosynthesis in these species. In the case of P. leucosticta, GP and ETR were additionally determined under saturating irradiance in algae pre-incubated for one week under white light at different irradiances and at white light (100 μmol m⁻² s⁻¹) enriched with far-red light. GP and growth rate increased at a growth irradiance of 500 μmol m⁻² s⁻¹ becoming photoinhibited at higher irradiances, while ETR increased when algae were exposed to the highest growth irradiance applied (2000 μmol m⁻² s⁻¹). The calculated O₂ evolved /ETR molar ratios were close to the theoretical value of 0.25 when algae were pre-incubated under 500–1000 μmol m⁻² s⁻¹. The enrichment by FR light provoked a decrease in both GP and ETR and an increase of nonphotochemical quenching although the irradiance of PAR was maintained at a constant level. In addition to C assimilation, other electron sinks, such as nitrogen assimilation, affected the GP–ETR relationship. The slopes of GP versus ETR or Φ_PSII versus Φ_{O_2} were lower in the algae with the highest N assimilation capacity, estimated as nitrate reductase activity and internal nitrogen contents, i.e., Ulva rotundata and Porphyra leucosticta, than that observed in U. olivascens. The possible mechanisms to explain this discrepancy between GP and ETR are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

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 acclimation to photon irradiance and its relation to chlorophyll fluorescence and carbon assimilation in the halotolerant green alga Dunaliella viridis

This work describes the long-term acclimation of the halotolerant microalga Dunaliella viridis to different photon irradiance, ranging from darkness to 1500 μmol m⁻² s⁻¹. In order to assess the effects of long-term photoinhibition, changes in oxygen production rate, pigment composition, xanthophyll cycle and in vivo chlorophyll fluorescence using the saturating pulse method were measured. Growth rate was maximal at intermediate irradiance (250 and 700 μmol m⁻² s⁻¹). The increase in growth irradiance from 700 to 1500 μmol m⁻² s⁻¹ did not lead to further significant changes in pigment composition or EPS, indicating saturation in the pigment response to high light. Changes in Photosystem II optimum quantum yield (F_v/F_m) evidenced photoinhibition at 700 and especially at 1500 μmol m⁻² s⁻¹. The relation between photosynthetic electron flow rate and photosyntetic O₂ evolution was linear for cultures in darkness shifting to curvilinear as growth irradiance increased, suggesting the interference of the energy dissipation processes in oxygen evolution. Carbon assimilation efficiencies were studied in relation to changes in growth rate, internal carbon and nitrogen composition, and organic carbon released to the external medium. All illuminated cultures showed a high capability to maintain a C:N ratio between 6 and 7. The percentage of organic carbon released to the external medium increased to its maximum under high irradiance (1500 μmol m⁻² s⁻¹). These results suggest that the release of organic carbon could act as a secondary dissipation process when the xanthophyll cycle is saturated. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthesis of lichen symbiotic alga Trebouxia erici as affected by irradiance and osmotic stress

The relation between oxygen evolution rate (OER) and quantum yield of photochemical reactions in photosystem 2 (Φ_PS2) was examined in lichen symbiotic alga Trebouxia erici Ahmadjian (strain UTEX 911) exposed to different irradiances and osmotic stress (2 M sucrose for 60 h). Linear relationship was found between OER and Φ_PS2 in control cell suspension within irradiance range of 0 – 500 μmol m⁻² s⁻¹. Under osmotic stress, OER and Φ_PS2 were significantly reduced. Relation between OER and Φ_PS2 was curvilinear due to strong osmotically-induced inhibition of OER at high irradiance. The highest used irradiance (500 μmol m⁻² s⁻¹) was photoinhibitory for osmotically-stressed T. erici because non-photochemical quenching (NPQ) increased substantially. Energy-dependent quenching represented major part of NPQ increase. Osmotic stress led also to the reduction of capacity of photochemical processes in PS 2 (F_V/F_M) and increase in F₀/F_M. These changes indicated negative effects of osmoticum on structure and function of photosynthetic apparatus.     

Characterization of acclimation of Hordeum vulgare to high irradiation based on different responses of photosynthetic activity and pigment composition

The ability of spring barley (Hordeum vulgare cv. Akcent) to adjust the composition and function of the photosynthetic apparatus to growth irradiances of 25–1200 μmol m⁻² s⁻¹ was studied by gas exchange and chlorophyll a fluorescence measurements and high-performance liquid chromatography. The increased growth irradiance stimulated light- and CO₂-saturated rates of CO₂ assimilation expressed on a leaf area basis up to 730 μmol m⁻² s⁻¹ (HL730), whereas at an irradiance of 1200 μmol m⁻² s⁻¹ (EHL1200) both rates decreased significantly. Further, the acclimation to EHL1200 was associated with an extremely high chlorophyll a/b ratio (3.97), a more than doubled xanthophyll cycle pool (VAZ) and a six-fold higher de-epoxidation state of the xanthophyll cycle pigments as compared to barley grown under 25 μmol m⁻² s⁻¹ (LL25). EHL1200 plants also exhibited a long-term inhibition of Photosystem II (PS II) photochemical efficiency (F _v/F _m). Photosynthetic capacity, chlorophyll a/b and VAZ revealed a linear trend of dependence on PS II excitation pressure in a certain range of growth irradiances (100–730 μmol m⁻² s⁻¹). The deviation from linearity of these relationships for EHL1200 barley is discussed. In addition, the role of increased VAZ and/or accumulation of zeaxanthin and antheraxanthin in acclimation of barley to high irradiance is studied with respect to regulation of non-radiative dissipation and/or photochemical efficiency within PS II. This revised version was published online in June 2006 with corrections to the Cover Date.     

Temperature and irradiance impacts on the growth,pigmentation and photosystem II quantum yields of <Emphasis Type="Italic">Haematococcus pluvialis</Emphasis> (Chlorophyceae)

The microalga Haematococcus pluvialis Flotow has been the subject of a number of studies concerned with maximizing astaxanthin production for use in animal feeds and for human consumption. Several of these studies have specifically attempted to ascertain the optimal temperature and irradiance combination for growth of H. pluvialis, but there has been a great deal of disagreement between laboratories. “Ideal” levels of temperature and irradiance have been reported to range from 14 to 28°C and 30 to 200 μmol photons m⁻² s⁻¹. The objective of the present study was to simultaneously explore temperature and irradiance effects for a single strain of H. pluvialis (UTEX 2505) across an experimental region that encompassed the reported “optimal” combinations of these factors for multiple strains. To this end, a two-dimensional experimental design based on response surface methodology (RSM) was created. Maximum growth rates for UTEX 2505 were achieved at 27°C and 260 μmol photons m⁻² s⁻¹, while maximum quantum yield for stable charge separation at PSII (F_v/F_m) was achieved at 27°C and 80 μmol photons m⁻² s⁻¹. Maximum pigment concentrations correlated closely with maximum F_v/F_m. Numeric optimization of growth rate and F_v/F_m produced an optimal combination of 27°C and 250 μmol photons m⁻² s⁻¹. Polynomial models of the various response surfaces were validated with multiple points and were found to be very useful for predicting several H. pluvialis UTEX 2505 responses across the entire two-dimensional experimental design space.     

Small scale vertical gradients of Arctic ice algal photophysiological properties

Photosynthetic parameters of phytoplankton and sea ice algae from landfast sea ice of the Chukchi Sea off Point Barrow, Alaska, were assessed in spring 2005 and winter through spring 2006 using Pulse Amplitude Modulated (PAM) fluorometry including estimates of maximum quantum efficiency (F _v/F _m), maximum relative electron transport rate (rETR_max), photosynthetic efficiency (α), and the photoadaptive index (E _k). The use of centrifuged brine samples allowed to document vertical gradients in ice algal acclimation with 5 cm vertical resolution for the first time. Bottom ice algae (0–5 cm from ice–water interface) expressed low F _v/F _m (0.331–0.426) and low α (0.098–0.130 (μmol photons m⁻²s⁻¹)⁻¹) in December. F _v/F _m and α increased in March and May (0.468–0.588 and 0.141–0.438 (μmol photons m⁻²s⁻¹)⁻¹, respectively) indicating increased photosynthetic activity. In addition, increases in rETR_max (3.3–16.4 a.u.) and E _k (20–88 μmol photons m⁻² s⁻¹) from December to May illustrates a higher potential for primary productivity as communities become better acclimated to under-ice light conditions. In conclusion, photosynthetic performance by ice algae (as assessed by PAM fluorometry) was tightly linked to sea ice salinity, temperature, and inorganic nutrient concentrations (mainly nitrogen).     

Irradiance prior to and during desiccation improves the tolerance to excess irradiance in the desiccated state of the old forest lichen Lobaria pulmonaria

Hydrated thalli of the lichen Lobaria pulmonaria were either preconditioned to dim irradiance (DI, 5 μmol m⁻² s⁻¹) or medium irradiance (MI, 200 μmol m⁻² s⁻¹) for 6 h. After this 6 h period, the thalli were allowed to desiccate under the two respective irradiances. Thereafter, these dry lichens were exposed to high irradiance (HI, 1 000 μmol m⁻² s⁻¹) for 60 h. After this HI treatment, the maximal photochemical quantum yield (F_V/F_M) and the de-epoxidation state of xanthophyll cycle pigments (DEPS) were highest in thalli preconditioned to MI. Hence irradiance in the last hydrated period before sampling is significant for the physiological state of lichens. A standardized irradiance pre-treatment before start of experiments is recommended.     

Photosynthetic and leaf anatomical characteristics of Castanea sativa: a comparison between in vitro and nursery plants

The anatomic and functional leaf characteristics related to photosynthetic performance of Castanea sativa growing in vitro and in nursery were compared. The irradiance saturated photosynthesis in in vitro grown plantlets was significantly lower compared to nursery plants (65 vs. 722 μmol m⁻² s⁻¹). The maximum photosynthetic rate (P_Nmax) was 4.0 and 10.0 μmol(CO₂) m⁻² s⁻¹ in in vitro microshoots and nursery plant leaves, respectively. Carboxylation efficiency (C_E) and electron transport rate (ETR) were three-folds higher in nursery plants than in microshoots. The nonphotochemical quenching (NPQ) was saturated at 80 μmol m⁻² s⁻¹ in microshoots suggesting limited photoprotection by thermal dissipation. The microshoots had wide open, spherical stomata and higher stomatal density than nursery plants and they had almost no epicuticular wax. Consequently, the microshoots had high stomatal conductance and high transpiration rate. These anatomic and functional leaf characteristics are likely major causes of the low survival rates of plantlets after ex vitro transfer.     

Effects of different irradiances on the photosynthetic process during ex-vitro acclimation of Anoectochilus plantlets

Six months old in vitro-grown Anoectochilus formosanus plantlets were transferred to ex-vitro acclimation under low irradiance, LI [60 μmol(photon) m⁻² s⁻¹], intermediate irradiance, II [180 μmol(photon) m⁻² s⁻¹], and high irradiance, HI [300 μmol(photon) m⁻² s⁻¹] for 30 d. Imposition of II led to a significant increase of chlorophyll (Chl) b content, rates of net photosynthesis (P _N) and transpiration (E), stomatal conductance (g _s), electron transfer rate (ETR), quantum yield of electron transport from water through photosystem 2 (Φ_PS2), and activity of ribulose-1,5-bisphosphate carboxylase/ oxygenase (RuBPCO, EC 4.1.1.39). This indicates that Anoectochilus was better acclimated at II compared to LI treatment. On the other hand, HI acclimation led to a significant reduction of Chl a and b, P _N, E, g _s, photochemical quenching, dark-adapted quantum efficiency of open PS2 centres (F_v/F_m), probability of an absorbed photon reaching an open PS2 reaction centre (F_v′/F_m′), ETR, Φ_PS2, and energy efficiency of CO₂ fixation (Φ_CO2/Φ_PS2). This indicates that HI treatment considerably exceeded the photo-protective capacity and Anoectochilus suffered HI induced damage to the photosynthetic apparatus. Imposition of HI significantly increased the contents of antheraxanthin and zeaxanthin (ZEA), non-photochemical quenching, and conversion of violaxanthin to ZEA. Thus Anoectochilus modifies its system to dissipate excess excitation energy and to protect the photosynthetic machinery.     

Physiological differences in the growth of <Emphasis Type="Italic">Sargassum horneri</Emphasis> between the germling and adult stages

The effects of temperature, irradiance, and daylength on Sargassum horneri growth were examined at the germling and adult stages to discern their physiological differences. Temperature–irradiance (10, 15, 20, 25, 30°C × 20, 40, 80 μmol photons m⁻²s⁻¹) and daylength (8, 12, 16, 24 h) experiments were carried out. The germlings and blades of S. horneri grew over a wide range of temperatures (10–25°C), irradiances (20–80 μmol photons m⁻²s⁻¹), and daylengths (8–24 h). At the optimal growth conditions, the relative growth rates (RGR) of the germlings were 21% day⁻¹ (25°C, 20 μmol photons m⁻²s⁻¹) and 13% day⁻¹ (8 h daylength). In contrast, the RGRs of the blade weights were 4% day⁻¹ (15°C, 20 μmol photons m⁻²s⁻¹) and 5% day⁻¹ (12 h daylength). Negative growth rates were found at 20 μmol photons m⁻²s⁻¹ of 20°C and 25°C treatments after 12 days. This phenomenon coincides with the necrosis of S. horneri blades in field populations. In conclusion, we found physiological differences between S. horneri germlings and adults with respect to daylength and temperature optima. The growth of S. horneri germlings could be enhanced at 25°C, 20 μmol photons m⁻²s⁻¹, and 8 h daylength for construction of Sargassum beds and restoration of barren areas.     

Eco-physiological responses of nitrogen-fixing cyanobacteria to light

The eco-physiological responses of three nitrogen-fixing cyanobacteria (N-fixing cyanobacteria), Aphanizomenon gracile, Anabaena minderi, and Ana. torques-reginae, to light were assessed under nutrient saturation. The N-fixing cyanobacteria were isolated into monocultures from a natural bloom in a shallow colored lake and their growth irradiance parameters and pigment composition were assessed. The different ecological traits related to light use (μ_max, α, I _k) suggest that these N-fixing cyanobacteria are well adapted to low light conditions at sufficient nutrients, yet interspecific differences were observed. Aphanizomenon gracile and Anabaena minderi had high relative growth rates at low irradiances (ca. 70% of those in high light), low half saturation constant for light-limited growth (I _k < 9.09 μmol photon m⁻² s⁻¹) and high efficiency (α < 0.11 day⁻¹ μmol photon⁻¹ m² s). Conversely, Ana. torques-reginae showed poorer light competitiveness: low relative growth rates at low irradiances (ca. 40% of those in high light), low α (0.009 day⁻¹ μmol photon⁻¹ m² s) and higher I _k (35.5 μmol photon m⁻² s⁻¹). Final densities in Aphanizomenon gracile and Anabaena minderi reached bloom densities at irradiances above 30 μmol photon m⁻² s⁻¹ with different hierarchy depending on irradiance, whereas Ana. torques-reginae never achieved bloom densities. All species had very low densities at irradiances ≤17 μmol photon m⁻² s⁻¹, thus no N-fixing blooms would be expected at these irradiances. Also, under prolonged darkness and at lowest irradiance (0 and 3 μmol photon m⁻² s⁻¹) akinetes were degraded, suggesting that in ecosystems with permanently dark sediments, the prevalence of N-fixing cyanobacteria should not be favored. All species displayed peaks of phycocyanin, but no phycoeritrin, probably due to the prevailing red light in the ecosystem from which they were isolated.     

Chlorophyll fluorescence in micropropagated <Emphasis Type="Italic">Rhododendron ponticum</Emphasis> subsp. <Emphasis Type="Italic">baeticum</Emphasis> plants in response to different irradiances

The aim of this study was to investigate acclimation of micropropagated plants of Rhododendron ponticum subsp. baeticum to different irradiances and recovery after exposure to high irradiance. Plants grown under high (HL) or intermediate (IL) irradiances displayed higher values of maximum electron transport rate (ETR_max) and light saturation coefficient (E_k) than plants grown under low irradiance (LL). The capacity of tolerance to photoinhibition (as assessed by the response of photochemical quenching, q_p) varied as follows: HL > IL > LL. Thermal energy dissipation (q_N) was also affected by growth irradiance, with higher saturating values being observed in HL plants. Light-response curves suggested a gradual replacement of q_p by q_N with increasing irradiance. Following exposure to irradiance higher than 1500 μmol m⁻² s⁻¹, a prolonged reduction of the maximal photochemical efficiency of PS 2 (F_v/F_m) was observed in LL plants, indicating the occurrence of chronic photoinhibition. In contrary, the decrease in F_v/F_m was quickly reverted in HL plants, pointing to a reversible photoinhibition.     

Excess irradiance causes early symptoms of senescence during leaf expansion in photoautotrophically <Emphasis Type="Italic">in vitro</Emphasis> grown tobacco plants

Photosynthetic parameters, growth, and pigment contents were determined during expansion of the fourth leaf of in vitro photoautotrophically cultured Nicotiana tabacum L. plants at three irradiances [photosynthetically active radiation (400–700 nm): low, LI 60 μmol m⁻² s⁻¹; middle, MI 180 μmol m⁻² s⁻¹; and high, HI 270 μmol m⁻² s⁻¹]. During leaf expansion, several symptoms usually accompanying leaf senescence appeared very early in HI and then in MI plants. Symptoms of senescence in developing leaves were: decreasing chlorophyll (Chl) a+b content and Chl a/b ratio, decreasing both maximum (F_V/F_M) and actual (Φ_PS2) photochemical efficiency of photosystem 2, and increasing non-photochemical quenching. Nevertheless, net photosynthetic oxygen evolution rate (P _N) did not decrease consistently with decrease in Chl content, but exhibited a typical ontogenetic course with gradual increase. P _N reached its maximum before full leaf expansion and then tended to decline. Thus excess irradiance during in vitro cultivation did not cause early start of leaf senescence, but impaired photosynthetic performance and Chl content in leaves and changed their typical ontogenetic course.     

Assessment of optimal depth and photon irradiance for cultivation of the brown alga, <Emphasis Type="Italic">Sargassum fulvellum</Emphasis> (Turner) C. Agardh

This study was aimed at determining the optimal depth and photon irradiance for growth of Sargassum fulvellum. Sampling and measurement of underwater irradiance were carried out at farms cultivating S. fulvellum at Wando, southwestern coast of Korea, from May 2004 to April 2005. Growth of thalli, underwater irradiance and photosynthetic quantum yield were measured over a range of depths for three culture stages. During their nursery cultivation stage (Stage I), length increase was greatest at 1.5 m depth (2.5 ± 0.2 cm), where the average midday irradiance over 28 days was 488 ± 58 μmol photons m⁻² s⁻¹. During the pre-main cultivation stage (Stage II), the greatest length increase occurred at 1 m depth (10.9 ± 0.1 cm) with an average irradiance of 845 ± 169 μmol photons m⁻² s⁻¹. For the main cultivation stage (Stage III) of the alga, thalli showed maximal length growth in March and early April at depths of 1–2 m and 3 m. These results suggest that growth at each cultivation stage of S. fulvellum could be controlled by depth of cultivation rope. Presented at the 6th Meeting of the Asian Pacific Society of Applied Phycology, Manila, Philippines.     

High irradiance increases organogenesis in friable callus of <Emphasis Type="Italic">Caustis blakei</Emphasis> Kük. (Cyperaceae)

Summary Caustis blakei is an attractive cut foliage plant harvested from the wild in Australia and marketed under the name of koala fern. Previous attempts to propagate large numbers of this plant have been unsuccessful. The effect of four light irradiances on organogenesis from compact and friable callus of C. blakei was studied for 21 wk. Both callus types produced numerous primordial shoots but many failed to develop into green plantlets. However, significantly more primordial shoots and green plantlets developed on the friable callus than on the compact callus, and significantly more green plantlets were regenerated under the higher photon irradiances of 200 and 300 μmol m⁻²s⁻¹ than under the lower irradiances of 100 and 150 μmol m⁻²s⁻¹. The compact callus produced its maximum number of green plantlets early in the experiment (after 9 wk), while the friable callus continued to produce primordial shoots and green plantelets throughout the period of the experiment, and reached its maximum production of green plantlets at 21 wk under the irradiance of 300 μmol m⁻²s⁻¹. Organogenesis from friable callus under high irradiance (300 μmol m⁻²s⁻¹) offers an efficient propagation method for C. blakei.     

Alterations in Rubisco activity and in stomatal behavior induce a daily rhythm in photosynthesis of aerial leaves in the amphibious-plant Nuphar lutea

Nuphar lutea is an amphibious plant with submerged and aerial foliage, which raises the question how do both leaf types perform photosynthetically in two different environments. We found that the aerial leaves function like terrestrial sun-leaves in that their photosynthetic capability was high and saturated under high irradiance (ca. 1,500 μmol photons m⁻² s⁻¹). We show that stomatal opening and Rubisco activity in these leaves co-limited photosynthesis at saturating irradiance fluctuating in a daily rhythm. In the morning, sunlight stimulated stomatal opening, Rubisco synthesis, and the neutralization of a night-accumulated Rubisco inhibitor. Consequently, the light-saturated quantum efficiency and rate of photosynthesis increased 10-fold by midday. During the afternoon, gradual closure of the stomata and a decrease in Rubisco content reduced the light-saturated photosynthetic rate. However, at limited irradiance, stomatal behavior and Rubisco content had only a marginal effect on the photosynthetic rate, which did not change during the day. In contrast to the aerial leaves, the photosynthesis rate of the submerged leaves, adapted to a shaded environment, was saturated under lower irradiance. The light-saturated quantum efficiency of these leaves was much lower and did not change during the day. Due to their low photosynthetic affinity for CO₂ (35 μM) and inability to utilize other inorganic carbon species, their photosynthetic rate at air-equilibrated water was CO₂-limited. These results reveal differences in the photosynthetic performance of the two types of Nuphar leaves and unravel how photosynthetic daily rhythm in the aerial leaves is controlled.     

Photosynthetic Response of Carrots to Varying Irradiances

Response to irradiance of leaf net photosynthetic rates (P _N) of four carrot cultivars: Cascade, Caro Choice (CC), Oranza, and Red Core Chantenay (RCC) were examined in a controlled environment. Gas exchange measurements were conducted at photosynthetic active radiation (PAR) from 100 to 1 000 μmol m⁻² s⁻¹ at 20 °C and 350 μmol (CO₂) mol⁻¹(air). The values of P _N were fitted to a rectangular hyperbolic nonlinear regression model. P _N for all cultivars increased similarly with increasing PAR but Cascade and Oranza generally had higher P _N than CC. None of the cultivars reached saturation at 1 000 μmol m⁻² s⁻¹. The predicted P _N at saturation (P _Nmax) for Cascade, CC, Oranza, and RCC were 19.78, 16.40, 19.79, and 18.11 μmol (CO₂) m⁻² s⁻¹, respectively. The compensation irradiance (I _c) occurred at 54 μmol m⁻² s⁻¹ for Cascade, 36 μmol m⁻² s⁻¹ for CC, 45 μmol m⁻² s⁻¹ for Oranza, and 25 μmol m⁻² s⁻¹ for RCC. The quantum yield among the cultivars ranged between 0.057–0.033 mol(CO₂) mol⁻¹(PAR) and did not differ. Dark respiration varied from 2.66 μmol m⁻² s⁻¹ for Cascade to 0.85 μmol m⁻² s⁻¹ for RCC. As P _N increased with PAR, intercellular CO₂ decreased in a non-linear manner. Increasing PAR increased stomatal conductance and transpiration rate to a peak between 600 and 800 μmol m⁻² s⁻¹ followed by a steep decline resulting in sharp increases in water use efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

The photosynthetic performance of the tropical seagrass Halophila ovalis in the upper intertidal

Some ecophysiological adaptation strategies of the tropical seagrass Halophila ovalis were investigated with respect to this plant's ability to grow in the upper intertidal in either monospecific pools (but not together with other intertidal species) or emergent and exposed to high temperatures and irradiances during several hours every day. It was found that Halophila ovalis could raise the pH in simulated pools to 8.6, while the two other major (biomass wise) intertidal seagrasses raised the pH to 8.8 (Cymodocea rotundata) and 9.2 (Thalassia hemprichii). In situ, midday pH values of 8.5, 8.7 and 9.0 were recorded in pools inhabiting the three species, respectively. It was further found that photosynthetic electron transport rates (ETR) measured continuously in situ for one leaf during a diurnal cycle followed the irradiance up to a daily maximal value of 2200 μmol photons m⁻² s⁻¹, but dropped by ca. 50% when the leaf became air exposed just before noon. This drop in ETR upon emergence was verified by “point measurements”, while ETRs of leaves that grew in small pools formed as the tide receded followed the irradiance more closely. Similarly, F_v/F_m measured after 15 min of dark adaptation decreased by ca. 50% in emergent leaves during midday, but maintained higher daily values in the submerged leaves.     

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.