Cell abundance and fluorescence of picoplankton in relation to growth irradiance and nitrogen availability in the red sea

The vertical distribution and cellular fluorescence characteristics (chlorophyll and phycoerythrin, PE) of picoplankton (cyanobacteria and prochlorophytes) in the southern Red Sea were investigated in relation to physico-chemical properties of the water column. At all stations two subpopulations of Synechococcus sp. (type A and B) co-occurred, with maximal numbers up to 75 000·cm⁻³. Type B, with dim fluorescence signals, dominated the surface waters whereas type A, with bright fluorescence signals, dominated at greater depth. The divinyl a and b containing Prochlorococcus sp. peaked below the cyanobacteria at the deep chlorophyll maximum (DCM) with maximal cell numbers of 276 000·cm⁻³.Due to thermal stratification the cellular fluorescence (chlorophyll and phycoerythrin) increased with decreasing growth (PAR) irradiance, in an S-shaped manner, but magnitude and slope for the three picoplankters differed. The Synechococcus sp. type B had only a moderate increase in chlorophyll and phycoerythrin fluorescence signals with depth (3.4 and 6.6 fold, respectively), with values saturating at 3% (L_d) of the surface irradiance. The deeper-water type not only possessed much higher values for cellular fluorescence than the B type, but the increase with decreasing light level was also much higer (for chlorophyll by a factor of 11 and PE increased by a factor of 23). In addition, maximal values for these fluorescence signals occurred at an isolume of 1 to 0.5%. These differences in concentrations and responses of the pigment content to the prevailing light climate explain the variation in abundance of both types over the water column.Although the prochlorophytes dominated almost the entire euphotic zone, their adaptation to low light levels was even better than in the two types of cyanobacteria. With depth their increase in chlorophyll fluorescence was similar to that observed in the cyanobacteria (with an increase from surface to bottom of the euphotic zone by a factor of 25). The maximum fluorescence signal was not reached until an I_d of 0.01%. The great ability of the prochlorophytes to adapt their pigmentation to changing light and nitrogen conditions suggests that they can maintain growth under a wide range of environmental conditions.     

Organic carbon in the upper 100 m and downward flux in the banda sea; monsoonal differences

Organic carbon was measured in the upper 100 m and its downward flux from the euphotic zone recorded, as part of a multidisciplinary study in the Banda Sea on monsoonal differences in the upper water layers.All 35 stations occupied showed on average 1.5 times higher POC values in August 1984 (4.9 g·m⁻², SE monsoon, upwelling period) than in February/March 1985 (3.3 g·m⁻², NW monsoon, downwelling period) for the 0 to 100 m water column. Monsoonal variation in POC was confined to the surface layer and related to a variation in phytoplankton content; it can be explained by — local— upwelling during the SE monsoon. At 100 m no seasonal variation occurred.Contrary to POC, average figures for DOC in February/March were almost twice the average observed in August (66.5 against 37.5 g·m⁻² for the 0 to 100 m water column). In February/March spatial variation was more pronounced than in August with highest values occurring in the NE part (>100 g·m⁻², 0 to 100 m), with a subsurface maximum. In this period the NW monsoon causes a surface current to the east. This might cause a westward subsurface countercurrent of Arafura Sea water rich in riverine or benthic layer DOC, influencing the eastern Banda Sea.At 100 m depth daily sedimentation amounted to ∼50 mg C·m⁻². The limited number of data do not indicate a difference between the 2 periods. On average the daily downward flux at 100 m depth amounted to 1.2% of POC and 0.8% of chlorophyll present in the 0 to 100 m water column, and equalled 5% of the daily primary production.     

Distribution of chroococcoid cyanobacteria and size-fractionated chlorophyll a biomass in the central and southern north sea waters during June/July 1989

The spatial and vertical distribution of phycoerythrin(PE)-containing chroococcoid cyanobacteria and the contribution of the <3 μm size fraction to overall phytoplankton chlorophyll a biomass were investigated in the central and southern North Sea during June and July 1989. PE-containing chroococcoid cyanobacteria cell numbers ranged between 6×10⁵ and 4.4×10⁷ cells·dm⁻³ and was typically between 0.5×10⁷ and 3×10⁷ cells·dm⁻³, lowest numbers being recorded near the British coast in central North Sea waters. The vertical distribution of these cyanobacteria showed no evidence to suggest a preferential accumulation deep in the euphotic zone. The <3 μm size fraction accounted for 6.6 to 57.5% of the total phytoplankton chlorophyll a biomass (mean 19.7%). In general, the relative significance of the <3 μm size fraction decreased with increasing total chlorophyll a biomass.     

Nutritional status and pigment composition of phytoplankton during spring and summer Phaeocystis blooms in Dutch coastal waters (Marsdiep area)

The general nutritional status and pigment composition of the phytoplankton population during the 1991 spring and summer Phaeocystis blooms in Dutch coastal waters was analysed using the 480/665 nm absorption ratio of 90% acetone extracts as well as reverse-phase HPLC-pigment analysis. A 100% increase in this ratio was indicative of moderate nutrient deficiency at the end of the Phaeocystis spring bloom in the Marsdiep area. During the summer, nitrate was more likely to be the limiting nutrient than phosphate. Both physiological adaptation and species composition were found to influence the absorption ratio. Variations in the absorption ratio did not completely reflect the variations in fucoxanthin and chlorophyll a. Therefore it was concluded that besides nutrient limitation dynamics, the variable abundances of in particular chlorophyll-c3 containing species (such as Prymnesiophyceae) may effect the absorption ratio.     

Characterization and mapping of novel chlorophyll deficient mutant genes in durum wheat

The yellow-green leaf mutant has a non-lethal chlorophyll-deficient mutation that can be exploited in photosynthesis and plant development research. A novel yellow-green mutant derived from Triticum durum var. Cappelli displays a yellow-green leaf color from the seedling stage to the mature stage. Examination of the mutant chloroplasts with transmission electron microscopy revealed that the shape of chloroplast changed, grana stacks in the stroma were highly variable in size and disorganized. The pigment content, including chlorophyll a, chlorophyll b, total chlorophyll and carotene, was decreased in the mutant. In contrast, the chla/chlb ratio of the mutants was increased in comparison with the normal green leaves. We also found a reduction in the photosynthetic rate, fluorescence kinetic parameters and yield-related agronomic traits of the mutant. A genetic analysis revealed that two nuclear recessive genes controlled the expression of this trait. The genes were designated ygld1 and ygld2. Two molecular markers co-segregated with these genes. ygld 1 co-segregated with the SSR marker wmc110 on chromosome 5AL and ygld 2 co-segregated with the SSR marker wmc28 on chromosome 5BL. These results will contribute to the gene cloning and the understanding of the mechanisms underlying chlorophyll metabolism and chloroplast development in wheat.     

Differential inheritance of pepper (Capsicum annuum) fruit pigments results in black to violet fruit colour

Colour is a critical determinant of fruit and vegetable quality. We characterized the inheritance of black and violet immature pepper fruit colour and chlorophyll, carotenoid and anthocyanin pigments responsible for fruit colour. Segregation for black vs. violet fruit colour deviated from expectations for simple inheritance. The primary determinant of fruit colour was the concentration of chlorophyll and carotenoid pigments relative to anthocyanin concentration. Whereas anthocyanin concentration was 34% greater in violet fruit, chlorophyll and carotenoid concentrations were 20‐fold and fourfold greater, respectively, in black fruit relative to concentrations found in violet fruit. Measurement of Commission Internationale d'Eclairage (CIE) colour space scores revealed larger a* values and smaller b* values for violet fruit relative to corresponding parameters for black fruit. An additive–dominance model was sufficient to explain the variation in individual fruit pigments as well as the CIE parameter a*. Addition of additive × additive effects to the additive–dominance model resulted in the best‐fit model m[d][h][i] for L* and b* colour space parameters. For all pigment classes, significant additive effects contributed to the genetic variance for fruit colour.     

Deep-water transit times in the eastern Indonesian basins,calculated from dissolved silica in deep and interstitial waters

The flushing of the deep basins in the eastern Indonesian archipelago is studied by means of the dissolved silica (Si) distribution, using water column data from the Snellius-II Expedition as well as from the Indopac Expedition. The linear θ-Si relationships below 500 to 1000 m permit the use of Si as a water-mass tracer. The main outflow of deep water from the Banda Sea into the Timor Trench is found east of Timor, above 1250 m depth, and the deeper parts of this outflow are partly recirculated through the Aru Basin and the Seram Sea into the north Banda Sea. Below the sill depth of the various basins, θ remains about constant while Si gradually increases towards the bottom. The time needed to create this Si-excess is obtained with the Si-flux out of sediments, calculated from pore-water gradients. Deep-water transit times are 20 years for the Banda Sea, 2 to 15 years for smaller basins and 60 years for the intermittently flushed Weber Deep. Modelling the Si-excess with error functions in some small basins gives high diffusion coefficients of 45 to 150 cm²·s⁻¹. The flux of Si from the sediments, 0.5 to 2 mol·m⁻²·a⁻¹, is higher than in other deep-sea basins, it amounts to roughly 30% of the biogenic silica production in the euphotic zone. Dissolved Si at 20–40 cm depth in the sediments ranges from 450 to 590 mmol·m⁻³; about 2% of Al in biogenic silica from sediments might cause this apparently low solubility for sediments rich in biogenic silica.     

Organic carbon in the water column and its sedimentation,fladen ground (North Sea), May 1983

POC, DOC and sedimentation were studied during the spring phytoplankton bloom from April 27 to May 25. POC in the euphotic layer consisted largely of phytoplankton, maximum values of 0.5 mg·dm⁻³ were found at the height of a bloom of the colonial microflagellate Corymbellus aureus. DOC increased during the waxing of this C. aureus bloom to a maximum of 2.4 mg·dm⁻³ after the bloom. Data indicate that most of the primary produced material ended up in the DOC pool. Sedimentation accounted only for small losses from the euphotic zone: ca 1% of the daily primary produced material sank out, except at the end of the C. aureus bloom when values increased to almost 10%. Daily sedimentation amounted to no more than 0.1 to 1% of the amounts present in the euphotic layer of POC, chlorophyll and phytoplankton cells.     

Physiological traits related to drought tolerance in tall fescue

The physiological basis of genetic variation in drought response and its association with yield and related indices is not clear in tall fescue. In this study thirty genotypes of tall fescue (Festuca arundinacea Schreb.) were sampled from a polycross population and evaluated under two levels of irrigation in 2010 (normal and intense stress) and 2011 (normal and mild stress). Physiological traits including relative water content (RWC), total chlorophyll (TChl), chlorophyll a (Chla), chlorophyll b (Chlb), Chla/Chlb, carotenoids (Car), TChl/Car and proline content along with forage yield, agro-morpholgical traits and selection indices (stress tolerance index, STI and drought susceptibility index, DSI) were studied. Large variation and moderate to high heritability was estimated for most of the studied traits. Intense drought condition decreased chlorophyll content while mild stress significantly increased it. In the other hand intense drought stress increased Chla/b while mild stress didn’t change it. Under mild drought stress condition STI was positively correlated with RWC while under intense drought stress condition STI was positively correlated with chlorophyll content. Although proline content was significantly increased in both intense and mild drought stress conditions, no relationship was found between proline accumulation with forage yield and STI. Applications of principle component analysis for screening suitable genotypes are also discussed.     

Identification and genetic mapping of a novel incompletely dominant yellow leaf color gene, <Emphasis Type="Italic">Y1718</Emphasis>, on chromosome 2BS in wheat

The yellow-green leaf color mutant (Ygm) is a spontaneous mutant derived from the common wheat (Triticum aestivum L.) cultivar Xinong1718. Genetic analysis has shown that a novel single incompletely dominant gene (Y1718) is responsible for the yellow leaf color phenotype. The progeny of Ygm exhibit three distinct leaf color phenotypes, i.e., yellow (Y), yellow-green (Yg), and normal green (G). Y plants have yellow-green leaves in the seedling stage, which become yellow or a strong gold-yellow in the booting stage, with dwarfism and thin tillers until the flowering stage, and underdeveloped thylakoid membranes without well-structured grana in the chloroplasts. Yg plants always have a yellow-green phenotype with a number of well-structured grana that are loosely connected with stroma lamellae in the chloroplasts, where their main agronomic traits are the same as Xinong1718 and G plants, but the seed yield is low. Compared with Xinong1718 and G plants, Y and Yg plants had much lower chlorophyll (Chl) a, Chl b, and carotenoid contents in the booting stage. Molecular analysis using an F₂ population and F_2:3 lines derived from a cross of Yg and Shannong1 indicated that the Y1718 gene is located on chromosome 2BS, where it is flanked by the simple sequence repeat marker Xwmc25 and expressed sequence tag-sequence tagged sites marker BE498358 at genetic distances of 1.7 and 4.0 cM, respectively. Our results facilitate the fine mapping and gene cloning of Y1718 to explore chlorophyll synthesis, metabolism, and development in wheat.     

Pheophytinase activity and gene expression of chlorophyll-degrading enzymes relating to UV-B treatment in postharvest broccoli (Brassica oleracea L. Italica Group) florets

Pheophytinase (PPH) activity and gene expression of chlorophyll (Chl)-degrading enzymes relating to UV-B treatment in postharvest broccoli (Brassica oleracea L. Italica Group) florets were determined. PPH is involved in the dephytylation of Mg-free Chl a, pheophytin (Phy) a. However, in vitro chlorophyllase (Chlase, EC.3.1.1.14) also uses Phy a as a substrate to produce pheophorbide (Pheide) a by dephytylation. For an accurate determination of PPH activity, the PPH protein fraction was separated from Chlase protein by ammonium sulfate precipitation. The protein precipitated by 45-60% saturated ammonium sulfate included a little bit of Chlase activity and was suitable for PPH determination. PPH activity in broccoli florets treated with a UV-B dose of 19 kJ m⁻² was repressed for the first 2 d of storage at 15 °C, whereas it increased gradually with senescence of control broccoli florets. The expression level of BoCLH1 was reduced in broccoli florets on day 4 of storage, while BoCLH2 and BoCLH3 were up-regulated with UV-B treatment. A high BoPAO expression level was found in senescent broccoli florets, and the up-regulation of this gene was delayed by UV-B treatment. The highest expression level of BoPPH was found in the control, and its expression was clearly repressed by UV-B treatment on day 2 of storage. We suggest that the up-regulation of Chl-degrading enzyme genes could be delayed by UV-B treatment, resulting in the suppression of floret yellowing in stored broccoli.     

Analysis of the main secondary metabolites produced in tomato (Lycopersicon esculentum,Mill.) epicarp tissue during fruit ripening using fluorescence techniques

Tomato (Lycopersicon esculentum L.) fruit are an important source of antioxidant (mainly pigment) compounds, as well as lycopene, β-carotene, ascorbic acid and polyphenols. Differentiation of the final product in the market requires an accurate evaluation of these value-adding compounds. Because of this, we have undertaken a comparison of the spectral characterisation of the tomato fruit surface pigments from the immature to over-ripe stage, using spectroscopy techniques based on visible fluorescence emission upon excitation in the same or ultraviolet spectral regions. The aim was to verify the spectral band for optimal conditions for fruit harvesting using non-destructive techniques. The pattern of pigment composition changed markedly during ripening and showed progressive disappearance of chlorophyll with a concomitant increase in carotenoids until the fully ripe stage. The main fluorescence spectral features belonging to anthocyanins, flavonoids, carotenoids and chlorophyll a after excitation of skin tomato pigments at different laser wavelengths was identified. In comparing, the fluorescence spectral ratios at the excitation wavelength λ_exc = 266 nm, significant differences were obtained for the spectral ratios of chlorophyll/flavonoids and carotenoids/chlorophyll. Positive correlation coefficients were found for the carotenoids/flavonoids (0.780) ratios and negative ones for the carotenoids/chlorophyll ratios (−0.513).Analysis of fluorescence resulted in determination of the most useful laser radiation for remote non-invasive measurements with laser-induced fluoresence (LIF): for the ripening stage, λ_exc = 266 nm was the optimal laser wavelength, since the induced fluorescence spectra obtained appeared to differ with the physiological stage of the fruit.     

The phytoplankton component of seston in San Francisco Bay

Phytoplankton biomass (as carbon) was estimated from chlorophyll a concentrations (Chla) and a mean value for the ratio of phytoplankton carbon to chlorophyll a in San Francisco Bay. The ratio was determined as the slope of a Model II regression of POC' against (Chla), where POC' is total particulate organic carbon minus sediment-associated non-phytoplankton carbon. Samples from 30 fixed sites in the channel and lateral shoals of San Francisco Bay were collected once or twice a month from April to November 1980, and at irregular intervals in South Bay during 1984 and 1985. For all data the calculated mean value of phytoplankton C:Chla was 51 (95% confidence interval = 47–54). No significant differences were found in the C:Chla ratio between shallow and deep sites (where light availability differs) or between northern and southern San Francisco Bay (where phytoplankton community composition differs). Using the mean C:Chla ratio of 51, we calculated that phytoplankton biomass constitutes about one third of seston carbon under most circumstances, but this fraction ranges from about 95% during phytoplankton blooms to less than 20% during spring periods of low phytoplankton biomass and high suspended sediment concentration.     

Distribution of photoperiod-insensitive alleles Ppd-B1a and Ppd-D1a and their effect on heading time in Japanese wheat cultivars

The genotypes of photoperiod response genes Ppd-B1 and Ppd-D1 in Japanese wheat cultivars were determined by a PCR-based method, and heading times were compared among genotypes. Most of the Japanese wheat cultivars, except those from the Hokkaido region, carried the photoperiod-insensitive allele Ppd-D1a, and heading was accelerated 10.3 days compared with the Ppd-D1b genotype. Early cultivars with Ppd-D1a may have been selected to avoid damage from preharvest rain. In the Hokkaido region, Ppd-D1a frequency was lower and heading date was late regardless of Ppd-D1 genotype, suggesting another genetic mechanism for late heading in Hokkaido cultivars. In this study, only 11 cultivars proved to carry Ppd-B1a, and all of them carried another photoperiod-insensitive allele, Ppd-D1a. The Ppd-B1a/Ppd-D1a genotype headed 6.7 days earlier than the Ppd-B1b/Ppd-D1a genotype, indicating a significant effect of Ppd-B1a in the genetic background with Ppd-D1a. Early-maturity breeding in Japan is believed to be accelerated by the introduction of the Ppd-B1a allele into medium-heading cultivars carrying Ppd-D1a. Pedigree analysis showed that Ppd-B1a in three extra-early commercial cultivars was inherited from ‘Shiroboro 21’ by early-heading Chugoku lines bred at the Chugoku Agriculture Experimental Station.     

Assessment of N topdressing rate at panicle initiation stage with chlorophyll meter-based diagnosis in rice

This study was conducted to determine N topdressing rate at panicle initiation stage by using chlorophyll meter (SPAD-502)-based N nutritional diagnosis in rice plant for high quality production of the Chucheongbyeo variety. Field experiments were carried for 3 years from 2003 to 2005 in Gyeonggi, Hwaseong. To confirm the method of measuring the leaf color with SPAD-502, leaf position of rice plant and measuring point on the leaf were studied. And the proper period to examine the leaf color and growth characteristics of rice plants was suggested at panicle initiation stage, from the results of the correlation between SPAD value and both content of chlorophyll and nitrogen in rice leaves. The multiply value of plant height, number of tillers per m² and SPAD value could explain well the effect of nitrogen fertilization on the growth of rice plants, and was used to determine an equation for on-site determination of nitrogen fertilization rate for Chucheongbyeo variety at the panicle initiation stage. Nitrogen fertilization increased number of tillers and rice yields, and affected the increase in protein content in rice grain resulting in loss of quality, such as taste value, and decreased the percentages of both ripened and whole rice grains. Based on the response surface statistical methodology using the SAS program, the relationship equation among protein content of brown rice (PROT), plant growth value (plant height × number of tillers × SPAD value, PGV), and nitrogen fertilization rate at the panicle formation stage (NF) was developed. That is, PROT (%) = 7.379403 ? (5.27E-7) a ? 0.013291 b + (3.025355E-13) a ² + (3.222997E-8) ab + (6.781E-05) b ², where a means PGV and b is NF (%), and the equation’s coefficient of determination (R²) was 0.967. From this equation, to recommend the optimum N fertilization rate at panicle initiation stage easily to produce high quality rice of Chucheongbyeo variety, which contains the target protein content of brown rice.     

Physical aspects of the flushing of the east Indonesian basins

The flushing of the deep basins in the east Indonesian archipelago is studied by means of water mass analysis, current observations and simple theoretical models. The Banda Sea and its secondary basins appear to be flushed from the Pacific Ocean with a typical flushing time of the order of 40 years. This fast flushing is made possible due to the high vertical diffusivity which is estimated to be of the order of 10⁻³ m²·s⁻¹. Theoretical models to descrive the vertical distribution of tracers due to mixing and basin wide upwelling turn out to fit the data very well. The observations cannot be used to confirm the horizontal structure of the deep circulation which results from a simple β-plane model. The vertical structure of the Aru Basin and the Timor Trench is more complicated due to the presence of water from the Indian Ocean, the Pacific Ocean and the Banda Sea. But also here simple models can be applied.     

Nuclear Mutations Affecting Chloroplastic Pigments of Photoperiod-insensitive Barley

Three photoperiod-sensitive spring barley cultivars (Hordeum vulgare L.) and three independently derived, single-gene, nuclear mutants expressing photoperiod insensitivity and extremely early heading time under short daylengths were investigated for chloroplastic pigment variation in three environments using reverse-phase high-performance liquid chromatography (HPLC) to account for differences in laminae colour. In a greenhouse, non-stress environment and in a full sunlight, high-temperature stress environment, no systematic differences were observed among pigments of the mutant-parent pairs. However, under 12 h of daily light (600 μE m ²sec^?1) and stress temperatures (20/10°C, night/day) in a growth chamber, the three mutants appeared similar to each other in chlorotic appearance and laminae pigment contents, but differed from the three non -chlorotic parents, which were similar to each other. The mutants had less chlorophyll a and b, β-carotene, lutein, taraxanthin, violaxanthin, and neoxanthin but more zeaxanthin than their parents. When shaded, the mutants became less chlorotic. How phenotypic differences for heading time and perception of day-length might be related to altered chloroplast contents remains unclear. The single-gene mutants conferring photopenod insensitivity were more sensitive to photothermal stress than their photoperiod-sensitive counterparts and as a result, their zeaxanthin content increased but the level of other pigments decreased.     

Single and Interactive Effects of Temperature and Light Quality on Four Canola Cultivars

Combined effects of temperature and light quality on plants have received little attention. We investigated the single and interactive effects of temperature and light quality on growth and physiological characteristics of four canola (Brassica napus) cultivars – Clearfield 46A76 (cv₁), Clearfield 45H72 (cv₂), Roundup Ready 45H24 (cv₃) and Roundup Ready 45H21 (cv₄). Plants were grown under lower (24°/20 °C) and higher (30°/26 °C) temperature regimes at low red/far‐red (R/FR), normal R/FR and high R/FR light ratios in environment‐controlled growth chambers (16 h light/8 h dark). Higher temperature reduced stem height and diameter; leaf number and area; dry matter of all plant parts; and specific leaf weight, but increased leaf area ratio; and chlorophyll (Chl) fluorescence (Y). Low R/FR increased stem height; Y; and ethylene, but decreased stem diameter; F_v/F_m; Chl a; Chl b; and carotenoids. Among cultivars, plants from cv₄ were tallest with thickest stems and greatest dry matter. None of the main factors affected gas exchange. Higher temperature at high R/FR caused cv₃ to be shortest, whereas lower temperature at low R/FR caused cv₄ to be tallest. We conclude that heat and other stress factors will adversely affect sensitive crops, but tolerant genotypes should perform well under future climate.     

Fine mapping and photosynthetic characteristics of the lower chlorophyll b 1 mutant in rice (Oryza sativa L.)

Chlorophylls absorb and transfer light energy to the photosynthetic system. Consequently, chlorophyll content is strongly related to crop biomass and yield. We isolated a rice spontaneous mutant, lower chlorophyll b 1 (lcb1), from a recombinant inbred line population. Under normal growth conditions, lcb1 plants produced yellow leaves with decreased total chlorophyll and chlorophyll b contents, but normal chlorophyll a content. Photosynthetic and fluorescence parameters differed between wild‐type and lcb1 plants. Compared with wild type, lcb1 had a higher electron transfer rate, a lower photochemical quenching coefficient and significantly reduced grain number, biomass and yield. A recessive nuclear gene controlled the mutant trait. Through map‐based cloning, we located the LCB1 gene in a 117.4‐kb region on the short arm of chromosome 3, close to the centromere, in a region containing 15 predicted candidate genes. None of these genes was directly related to chlorophylls or the chloroplast; therefore, lcb1 may be a mutation of a novel gene. These results will be useful for further research on the molecular mechanisms controlling biogenesis and chloroplast biochemical processes.