Water cleavage by solar radiation—an inspiring challenge of photosynthesis research

Solar energy exploitation by photosynthetic water cleavage is of central relevance for the development and sustenance of all higher forms of living matter in the biosphere. The key steps of this process take place within an integral protein complex referred to as Photosystem II (PS II) which is anisotropically incorporated into the thylakoid membrane. This minireview concentrates on mechanistic questions related to i) the generation of strongly oxidizing equivalents (holes) at a special chlorophyll a complex (designated as P680) and ii) the cooperative reaction of four holes with two water molecules at a manganese containing unit WOC (water oxidizing complex) resulting in the release of molecular oxygen and four protons. The classical work of Pierre Joliot and Bessel Kok and their coworkers revealed that water oxidation occurs via a sequence of univalent oxidation steps including intermediary redox states S_i (i = number of accumulated holes within the WOC). Based on our current stage of knowledge, an attempt is made a) to identify the nature of the redox states S_i, b) to describe the structural arrangement of the (four) manganese centers and their presumed coordination and ligation within the protein matrix, and c) to propose a mechanism of photosynthetic water oxidation with special emphasis on the key step, i.e. oxygen-oxygen bond formation. It is assumed that there exists a dynamic equilibrium in S₃ with one state attaining the nuclear geometry and electronic configuration of a complexed peroxide. This state is postulated to undergo direct oxidation to complexed dioxygen by univalent electron abstraction with Y_Z ^ox and simultaneous internal ligand to metal charge transfer.Key questions on the mechanism will be raised. The still fragmentary answers to these questions not only reflect our limited knowledge but also illustrate the challenges for future research.Abbreviations b559 cytochrome b559 - BChl bacteriochlorophyll - Chl chlorophyll - CP47 Chl a containing a 47 kDa polypeptide - D1/D2 polypeptides of the PS II reaction center - ENDOR electron nuclear double resonance - EPR electron paramagnetic resonance - ESEEM electron spin echo envelope modulation - EXAFS extended X-ray absorption fine structure - FTIR Fourier transform infrared - NMR nuclear magnetic resonance - P680, P700 photoactive Chl a of PS II and PS I, respectively - PS II Photosystem II - Q_A special plastoquinone of PS II - S_i redox states of WOC - WOC water oxidizing complex - WOS water oxidizing site - UV/VIS ultraviolet/visible - Y_D, Y_Z redox active tyrosines of polypeptides D2 and D1, respectively     

Thylakoid Membrane Development and Differentiation: Assembly of the Chlorophyll a-b Light-Harvesting Complex and Evidence for the Origin of Mr=19, 17.5 and 13.4 kDa Proteins

Pea plants were grown under intermittent illumination (ImL)conditions. The low dosage of light given to ImL plastids limitedthe rate of chlorophyll (Chl) a and Chl b biosynthesis and,therefore, it retarded the rate of photosynthetic unit formationand thylakoid membrane development. Depending on the developmentalstage of the photosynthetic unit, ImL plastids had variableChl a/Chl b ratios (2.7 <Chl a/Chlb<20) and showed distinctintermediates in the assembly of the chlorophyll a–b light-harvestingcomplex (LHC) of photosystem-II (PSII). The results are consistentwith a step-wise increment in the PSII antenna size involvingthree distinct forms of the PSII unit: (i) a PSII-core formwith about 37 Chl a molecules; (ii) a PSIL_ß form containingthe PSII-core and the LHC-II-inner antenna with a total of about130 Chl (a + b) molecules, and (iii) the mature PSII_a form containingPSII_ß and the LHC-II-peripheral antenna with a totalof 210–300 Chl (a + b) molecules. The thylakoid membranecontained polypeptide subunits b, c and d (the Lhcb1, 2 and3 gene products, respectively) when only the LHC-II-inner waspresent. Polypeptide subunit a, (the apoprotein of the chlorophyll-proteinknown as CP29), along with increased amounts of b and c appearedlater in the development of thylakoids, concomitant with theassembly of the LHC-II-peripheral. The results suggest thatpolypeptide subunit d has priority of assembly over subunita. It is implied that, of all LHC-II constituent proteins, subunitd is most proximal to the PSII-core complex and that it servesas a linker in the transfer of excitation energy from the bulkLHC-II (subunits b and c) to the PSII-core. The work also addressesthe origin of low-molecular-weight proteins (M_r = 19, 17.5 and13.4 kDa) which co-isolate with intact developing plastids andwhose abundance decreases during plastid development. Aminoacid compositional and immunoblot analyses show a nuclear histoneorigin for these low-molecular-weight proteins and suggest co-isolationof histone-containing nuclear vesicles along with intact developingplastids. ¹Present address: Plant Physiology Research Group, The Universityof Calgary, Department of Biological Sciences, 2500 UniversityDrive N.W., Calgary, Alberta CANADA T2N 1N4.     

Temporal and vertical variation of chlorophyll a concentration, phytoplankton photosynthetic activity and light attenuation in Lake Kinneret: possibilities and limitations for simulation by remote sensing

The relationship between chlorophyll a (Chl a) and primary productivity(PP) in the uppermost water layer and the water column-based(0–15 m) integral values of those variables were examinedusing measurements taken in Lake Kinneret (Israel) from 1990to 2003. In 81% of all Chl a profiles examined, the distributionwas fairly uniform within the entire 0–15 m water column,and 12.3% of instances showed a prominent subsurface maximum,when the lake phytoplankton was dominated by the dinoflagellatePeridinium gatunense. Chl a can be reliably estimated by remotesensing techniques in the productive and turbid water of LakeKinneret, since Chl a concentration at surface layers can beextrapolated to the entire water column. Light vertical attenuationcoefficient average for wavelengths from 400 to 700 nm, K_d,ranged from 0.203 to 1.954 m^–1 and showed high degreeof temporal variation. The maximal rate of photosynthetic efficiency,P_B^opt [average 3.16 (±1.50)], ranged from 0.25 to 8.85mg C m^–3 h^–1 mg Chl a^–1. Using measured dataof Chl a, P_B^opt, and light as an input, a simple depth-integratedPP model allowed plausible simulation of PP. However, a lackof correlation between photosynthetic activity and temperature(or other variable with remotely sensed potential) renders theuse of models that require input of photosynthetic efficiencyto calculate integrated PP of little value in the case of productiveand turbid Lake Kinneret.     

Biogeographic study of the planktonic communities of the Prince Edward Islands (Southern Ocean)

Microphytoplankton and zooplankton composition and distributionin the vicinity of the Prince Edward Islands and at the Sub-antarcticFront (SAF) were investigated in late austral summer (April/May)1996. Samples were collected for analysis of chlorophyll a concentration(Chi a), microphytoplankton and zooplankton abundance. Generally,the highest Chl a concentrations (up to 2.0 µg l^–1)and zooplankton densities (up to 192 ind. m^–3) were recordedat stations within the inter-island area while the lowest values(<0.4 µg l^–1) were observed at stations upstreamof the islands. High Chl a and zooplankton biomass values werealso associated with the SAF. Microphytoplankton were dominatedby chain-forming species of the genera Chaetoceros (mainly C.neglectus),Fragilariopsis spp. and the large diatom Dactyliosolen antarcticus.The zooplankton assemblages were always dominated by mesozooplanktonwhich at times contributed up to 98% of total zooplankton abundanceand up to 95% of total biomass. Among mesozooplankton, copepods,mainly Clausocalanus brevipes and Metridia lucens numericallydominated. Among the macrozooplankton euphausiids, mainly Euphausiavallentini, E.longirostis and Stylocheiron maximum, and chaetognaths(Sagitta gazellae) accounted for the bulk of abundance and biomass.Cluster and ordination analysis did not identify any distinctbiogeographic regions among either the microphytoplankton orzooplankton.     

Phytotoxicity of Phthalate Plasticisers: 2. SITE AND MODE OF ACTION

The effects of phthalate esters on chlorophyll a₂ fluorescencein radish plants (Raphanus sativus L. cv. Cherry Belle) wereexamined Fluorescence yield was increased in those plants exposedto an aerial concentration of 120 ng dm^–3 dibutyl phthaiatc(DBP) at a rate of 3.0 dm³ min^–1 for 13 d. Comparisonof fluorescence enhancement ratios and F_red/F_ox, suggests thatDBP inhibits photosynthesis in radish plants at a site afterQ_A. Both DBP and diisobutyl phthalate (DIBP) strongly inhibiteduncoupled (PS2+PS1) electron transport rates in thylakoids isolatedfrom spinach. At a chlorophyll concentration of 10 µgcm^–3 the concentrations of DBP and DIBP exhibiting 50%inhibition were 44 mmol m^–3 and 42 mmol m^–3 respectively.Basal electron transport rates were also inhibited, with 87mmol m^–3 of DBP or DIBP producing 50% inhibition. Measurementof photosystcm 1 activity suggested that the main site of actionof these phthalates was localized at a site near the reducingside of photosystem 2. Key words: Phthalate, plasticiser, chlorophyll, fluorescence, photosynthesis, inhibition     

Trophic structure of nine Czech reservoirs regularly stocked with piscivorous fish

For nearly 20 years, most Czech reservoirs supplying drinking water have been under statutory protection which permitted reservoir managers to manipulate fish stocks in order to maintain a sustainable water quality. The most common biomanipulative measure adopted was stocking with piscivorous fish (mostly 5 cm fry) using an annual stocking level of approx. 25 000 fish per reservoir. Nine reservoirs were studied for signs of top-down food web effects, as predicted by the trophic cascade hypothesis based on levels of total phosphorus (TP), chlorophyll a (Chl a), zooplankton biomass (ZB) and zooplankton community structure. In all nine reservoirs, only small Daphnia species were recorded, such as D. galeata and D. cucullata. The proportion of large-bodied daphnids retained on a 0.71 mm sieve was less than 10% of the total crustacean biomass in all reservoirs. The relationship of Chl a level – TP, and of ZB – Chl a, was positive under enhanced piscivory and did not differ statistically from the relationships in other reservoirs with natural fish stocks. This implies that bottom-up forces remained stronger than top-down ones in the studied reservoirs, despite the stocking of piscivorous fish. The failure of this attempt at biomanipulation may be due to an insufficient stocking rate of predatory fish and/or inadequate data on the resident planktivorous fish levels.     

On the relationship between carbon fixation efficiency and bio-optical characteristics of phytoplankton

Bio-optical characteristics of phytoplankton and carbon assimilationwere measured in the NW Atlantic Ocean near the continentalshelf break South of New England (40°N/70°W) in August2002. Discrete samples included: chlorophyll a (Chl a), phytoplanktonabsorption (a_ph), the photosynthetic versus irradiance (P versusE) response using carbon fixation, the maximum quantum yieldfor charge separation at Photosystem II (PSII) (F_v/F_m) and itsrecovery kinetics and the functional absorption cross-section(_PSII). In this study, we present a robust empirical relationshipthat illustrates the possibility of estimating photosyntheticcarbon uptake using independent measures of absorption and variablefluorescence. Covariations between the initial slope of thephotosynthesis-irradiance curve normalized per Chl a, the a_phindex of the ratio of photoprotective to total photosyntheticcarotenoids and the dark recovery of the maximum quantum yieldefficiency of PSII suggest a new pathway to estimate primaryproduction in the marine environment, including the effect ofthe recent light history. Recovery kinetics in laboratory cultureconfirmed the effects of light history in the derived parameterfrom the dark incubations from natural populations.     

Properties of Chlamydomonas Photosystem II Core Complex with a His-Tag at the C-Terminus of the D2 Protein

A His-tagged PSII core complex was purified from recombinantChlamydomonas reinhardtii D2-H thylakoids by single-step Ni²⁺-affinitycolumn chromatography and its properties were partially characterizedin terms of their PSII functions and chemical compositions.The PSII core complex that has a His-tag extension at the C-terminusof the D2 protein evolved oxygen at a high rate of 2,400 µmol(mg Chl)^–1h^–1 at the optimum pH of 6.5 with ferricyanideand 2,6-dichlorobenzoquinone as electron acceptors in the presenceof Ca²⁺ as an essential cofactor, and approximately 90% of theactivity was blocked by 10 µM DCMU. The core complex exhibitedthe thermoluminescence Q-band but not the B-band regardlessof the presence or absence of DCMU, although both bands wereobserved in the His-tagged thylakoids. The core complex wasfree from PSI and contained one Y_D, Tyr 160 of the D2 protein,four Mn atoms, two cytochrome b-559, about 46 Chl a molecules,and probably one Q_A, the primary acceptor quinone of PSII. Itwas inferred from these results that His-tagging at the C-terminusof the D2 protein does not affect the functional and structuralintegrity of the PSII core complex, and that the ‘His-tagstrategy’ is highly useful for biochemical, physicochemical,and structural studies of Chlamydomonas PSII. (Received October 22, 1998; Accepted December 25, 1998)     

Relative effects of Daphnia and Ceriodaphnia on phosphorus-chlorophyll relationships in small urban lakes

Empirical models based on zooplankton biomass were used to predict mean summer chlorophyll a (Chl a) and to examine how zooplankton influenced the total phosphorus (TP) - Chl a relationship. Four years of data were analyzed for three lakes having similar TP concentrations but varied abundances of Daphnia and Ceriodaphnia. Mean TP did not correlate significantly with mean Chl a during the study period, although mean Daphnia density was a good predictor of Chl a concentration (p > 0.001). Both residuals from the TP - Chl a relationship (p > 0.001) and Secchi depth (p > 0.007) were negatively correlated with Daphnia abundance. Ceriodaphnia abundance was positively correlated with Chl a (p > 0.002) and Secchi depth (p > 0.001). Mean size of Daphnia during spring was the best predictor of the Daphnia-Ceriodaphnia shift in mid-summer. Early establishment of a large-sized Daphnia cohort may prevent their summer elimination by Chaoborus and intensify competition with Ceriodaphnia. These results imply an important link between Daphnia and Ceriodaphnia thereby limiting the utility of Chl a - TP model predictions in these small, urban lakes. This linkage and the differential effect of these two zooplankton species on planktonic algae deserve further consideration in similar lakes where phytoplankton and zooplankton tend to be tightly coupled.     

Growth and grazing losses of prokaryotes in the central Atlantic Ocean

The trophic relation between prokaryotes and heterotrophic nanoflagellateswas studied during two latitudinal cruises in the central AtlanticOcean. The losses to predation on prokaryotes were determinedin 12 locations covering a wide range of trophic situations,from ultraoligotrophic [<0.05 mg chlorophyll a (Chl a) m^–3]to moderately eutrophic waters (>1 mg Chl a m^–3). Inthese locations, the abundance of prokaryotes (P) covaries withthat of heterotrophic nanoflagellates, thus suggesting thatresources controlled the abundance of heterotrophic nanoflagellates(HNF). Besides, the losses to predation were positively relatedto prokaryotic and heterotrophic nanoflagellate biomass, whichpoints toward higher consumption rates associated with largerconcentrations of preys and predators. Conversely, decliningtrends between prokaryotic production (P_P) and the fractionof this production lost to predation revealed higher relativelosses in the environments with lower productions. Our studyshows for the central Atlantic that 35% of prokaryotic biomass(B_P), equating to between 40 and 83% of P_P can be ingested dailyand that 55% of the variability observed in the rate of prokaryoticloss to predation was related with the HNF. As predators grazeon many prey types, in an oligotrophic system containing manyprey species but little numeric loading, there will still beprey for predators but not enough hosts for viruses. In thissense, our study confirms the importance of the prey–predatorrelationship between prokaryotes and heterotrophic nanoflagellatesin the flow of carbon of the less productive regions of theocean.     

外源NO对增补UV-B辐射下兴安落叶松幼苗叶片光合色素和叶绿素荧光特性的影响

在增强UV-B辐射下,以3年生兴安落叶松幼苗为实验材料,研究了外源NO供体硝普钠(Sodium nitroprusside,SNP)对幼苗的光合色素（Chla、Chlb和Car）和叶绿素荧光参数的影响。方差分析结果表明0.5 mmol·L^-1的SNP对增补UV B胁迫下的兴安落叶松幼苗产生显著影响。0.5 mmol·L^-1的SNP能够显著抑制增补UV-B辐射后光合色素、F_v/F_m、Φ_PSⅡ、F_v′/F_m′和qP的明显下降以及Chla /Chlb、F_o和NPQ的升高。表明了外源NO能够减轻UV-B辐射胁迫下兴安落叶松幼苗光合反应中心的生理损伤,从而增强兴安落叶松幼苗对增补UV-B辐射胁迫环境的适应能力。     

Seasonal and interannual variability of chlorophyll a and primary production in the Equatorial Atlantic: in situ and remote sensing observations

The seasonal variability of phytoplankton in the EquatorialAtlantic was analysed using Sea-viewing Wide Field-of-view Sensor(SeaWiFS)-derived chlorophyll a (Chl a) concentration data from1998 to 2001, together with in situ Chl a and primary productiondata obtained during seven cruises carried out between 1995and 2000. Monthly averaged SeaWiFS Chl a distributions werein agreement with previous observations in the Equatorial Atlantic,showing marked differences between 10° W in the EasternTropical Atlantic (ETRA) and 25° W in the Western TropicalAtlantic (WTRA) provinces (Longhurst et al. 1995. J. PlanktonRes., 17, 1245–1271). The seasonal cycle of SeaWiFS-derivedChl a concentration calculated for 0–10° S, 0–20°W (ETRA) is consistent with in situ Chl a measurements, withvalues ranging from 0.16 mg m^–3, from February to April,to 0.52 mg m^–3 in August. Lower variability was observedin 10° N–10° S, 20–30° W (WTRA) whereminimum and maximum concentrations occurred in April (0.15 mgm^–3) and in August (0.24 mg m^–3), respectively.A significant empirical relationship between depth-integratedprimary production and in situ measured sea surface Chl a wasfound for ETRA, allowing us to estimate the seasonal cycle ofdepth-integrated primary production from SeaWiFS-derived Chla. As for Chl a, this model was verified in a small area ofthe Eastern Equatorial Atlantic (0–10° S, 0–20°W), although in this instance it was not completely able todescribe the magnitude and temporal variability of in situ primaryproduction measurements. The annual euphotic depth-integratedprimary production rate estimated for ETRA by our empiricalmodel was 1.4 Gt C year^–1, which represents 16% of theopen ocean primary production estimated for the whole AtlanticOcean.     

Respiratory ETS activity of plankton in the northwestern Alboran Sea: seasonal variability and relationship with hydrological and biological features

Respiratory electron transport system (ETS) activity was measuredin plankton samples (<200 µm) collected in the NW AlboranSea. Sampling was carried out during seasonal cruises (summerand autumn 2003 and winter and spring 2004) in 12 stations locatedin transects off the coast of Malaga (southern Spain). Thiswork reports for the first time seasonal variations of the Arrheniusactivation energy (Ea) as well as being the first study to addressCO₂ balance in the NW Alboran Sea. These variations were relatedto changes in the phytoplankton community assemblage, whichcould ultimately be caused by the seasonal variability of hydrologicalconditions. ETS activity was significantly higher in summer,coinciding with a higher chlorophyll a (Chl a) concentrationand relatively high levels of particulate organic matter. TheETS:Chl a_total ratios were low during the four seasons, suggestinga high contribution of autotrophic phytoplankton to the respiratoryactivity of planktonic community. Respiratory CO₂ production(RCP) calculated from ETS activity ranged from 4.6 to 28.1 mgC m^–3 day^–1 during the four cruises. Chl a-specificRCP was lower than the maximum photosynthetic rates reportedin the literature for the studied area, suggesting that primaryproduction (PP) and respiration in the water column might beunbalanced.     

Chromatographic separation of photosystems I and II from the thylakoid membrane isolated from a thermophilic blue-green alga

The thylakoid membrane of a thermophilic blue-green alga, Synechococcussp., was separated into four chlorophyll-containing fractionsby a single chromatographic manipulation with a diethylaminoethyl-cellulosecolumn after digitonin treatment. Photosystems I and II, orchlorophyll a forms, were unevenly distributed among the fourfractions, which were designated F-1, F-2, F-3 and F-4 in theorder of elution from the column. F-1 has a simple composition of the chlorophyll a form and totallylacks photochemical activity. This fraction may be an antennachlorophyll a-protein in the blue-green alga. F-2 is rich inshorter wavelength chlorophyll a forms and shows the three-bandedfluorescence emission spectrum characteristic of photosystemII at liquid nitrogen temperature. This fraction is highly activein 2,6-dichloroindophenol photoreduction and contains one photooxidizablecytochrome b₅₅₉ per 50–100 chlorophyll a, whereas theP-700 content is as low as one P-700 per 2,000 chlorophyll a.Thus, F-2 represents photosystem II in a highly purified state.F-3 is rich in photosystem I, since this fraction is inactivein 2,6-dichloroindophenol photoreduction, and contains one P-700per 200 chlorophyll a and smaller amounts of cytochrome b₅₅₉.Longer wavelength chlorophyll a forms are abundant and a peakat 730 nm is the most prominent in the low-temperature fluorescencespectrum in this fraction. F-4, which consists of larger membranefragments shows spectral and photochemical features similarto those of F-3. (Received August 8, 1979; )     

Chromatic Regulation of Photosystem Composition in the Photosynthetic System of Red and Blue-Green Algae

A chromatic adaptation in the photosynthetic quantum yield forthe light mainly absorbed by chlorophyll a (Chl a light) firstfound by Yocum (1951) was studied with one red and three blue-greenalgal strains. When the cells were grown under a weak Chl alight, the quantum yield in all the strains increased. Comparisonof photosystem (PS) compositions, including phycobilin (PBP)and Chl a antennae, reaction centers I and II, in the cellsgrown under the light mainly absorbed by PBP and Chl a revealedthat changes in quantum yield could be attributed to changesin the ratio of PS I/II; PS I/II becomes larger than 1 underPBP light but decreases to 1 in most cases under Chl a light.The change in the PS I/II ratio is due solely to the changesin the PS I population in the cell; PS II remains constant.These results are similar to the intensity-dependent responsein PS composition. A common hypothesis for both the chromatic and intensity-inducedregulation of PS composition was proposed based on the ideaof balance between the electron flow from H₂O to NADP drivenby PS I and II and the cyclic one driven by PS I. (Received May 16, 1985; Accepted September 4, 1985)     

Extinction coefficients of chlorophyll a and B in n,n-dimethylformamide and 80% acetone

We found inconsistencies in the commonly used data for chlorophyll analysis in 80% acetone. Recently developed extinction coefficients for chlorophyll b in N,N-dimethylformamide (DMF) based on values from 80% acetone are low as a result of these inconsistencies. We determined extinction coefficients of chlorophyll a (Chl a) and chlorophyll b (Chl b) in DMF for wavelengths of 618 to 665 nanometers. The simultaneous equations necessary for quantifying Chl a, Chl b, or total Chl in DMF in the absence of other chlorophyllous pigments are: Chl a = 12.70A_664.5 - 2.79A₆₄₇; Chl b = 20.70 A₆₄₇ - 4.62A_664.5; total Chl = 17.90A₆₄₇ + 8.08A_664.5, where A = absorbance in 1.00 centimeter cuvettes and Chl = milligrams per liter.

N,N-Dimethylformamide is a very convenient solvent for Chl extraction since it is effective on intact plant parts and Chl is quite stable in DMF. There was no difference in the amount of Chl extracted when plant tissue was stored for 1 or 3 days at three temperatures, with or without solvent added.

     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

Zooplankton in the Ligurian Sea: Part II. Exploration of their physical and biological forcing functions during summer 2000

A survey of the biological and physical oceanography of theLigurian Sea was conducted in the late summer of 2000. Forty-onestations were sampled for nutrients, oxygen, fluorescence andhydrographic information. Acoustic backscatter measurementswere used to estimate abundance of small (<5 mm) zooplanktonbiovolume versus depth and the distribution of northern krill,Meganyctiphanes norvegica. Net-tow and underwater video datawere collected to identify the zooplankton present. These datawere used to analyze the Ligurian Sea ecosystem for physicaland biological linkages that control zooplankton abundance anddistribution. Results are compared with those from a similarstudy conducted in 1999. Hydrographic sampling showed a domeof dense water in the southwestern middle of the basin. Thehighest chlorophyll a (Chl a) concentrations were measured inthis area, while small zooplankton biovolume was evenly distributedthroughout the survey. Integrated values of Chl a and smallzooplankton biovolume in 2000 were greater than in 1999. Meganyctiphanesnorvegica, siphonophores and salps were the dominant componentsof the macrozooplankton population in the upper 200 m. In thesampled depth strata, siphonophore abundance did not changeduring the day, while M. norvegica were only caught at night.Acoustic backscatter data show that higher densities of M. norvegicaoccurred in deeper water and in the western and southwesternareas of the Ligurian Sea.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.