Genetic analysis of biomass and photosynthetic parameters in wheat grown in different light intensities

Growth light intensities largely determine photosynthesis, biomass, and grain yield of cereal crops. To explore the genetic basis of light responses of biomass and Researchphotosynthetic parameters in wheat(Triticum aestivum L.),a quantitative trait locus(QTL) analysis was carried out in a doubled haploid(DH) population grown in low light(LL),medium light(ML), and high light(HL), respectively. The results showed that the wheat seedlings grown in HL produced more biomass with lower total chlorophyll content(Chl), carotenoid content, and maximum photochemical efficiency of photosystem II(Fv/Fm) while the wheat seedlings grown in LL produced less biomass with higher Chl compared with those grown in ML. In total, 48 QTLs were identified to be associated with the investigated parameters in relation to growth light intensities. These QTLs were mapped to 15 chromosomes which individually explained6.3%–36.0% of the phenotypic variance, of which chromo-somes 3A, 1D, and 6B were specifically involved in LL response, 5D and 7A specifically involved in ML response,and 4B specifically involved in HL response. Several light-responsive QTLs were co-located with QTLs for photosynthetic parameters, biomass, and grain weight under various conditions which may provide new hints to uncover the genetic control of photosynthesis, biomass, and grain weight.     

Effect of copper deficiency on photosynthetic electron transport in wheat plants

Copper deficiency in wheat ( Triticum aestivum L. cv. Nazareno Stramppeli) markedly affects photosynthetic activity. Flag leaves of copper-deficient plants showed a 50% reduction of the photosynthetic rate expressed as mg CO₂ dm⁻²h⁻¹. The activities of PSI and PSII, determined for isolated chloroplasts, as well as fluorescence measurements on intact leaves of copper-deficient plants, indicated a low activity of photosynthetic electron transport. Ribulose bisphosphate carboxylase/oxygenase (Rubisco) activity was not affected by copper deficiency but copper deficiency affected the chloroplast ultrastructure, especially at the level of grana, where a disorganization of thylakoids is evident.     

Antioxidant responses of wheat plants under stress

Currently, food security depends on the increased production of cereals such as wheat (Triticum aestivum L.), which is an important source of calories and protein for humans. However, cells of the crop have suffered from the accumulation of reactive oxygen species (ROS), which can cause severe oxidative damage to the plants, due to environmental stresses. ROS are toxic molecules found in various subcellular compartments. The equilibrium between the production and detoxification of ROS is sustained by enzymatic and nonenzymatic antioxidants. In the present review, we offer a brief summary of antioxidant defense and hydrogen peroxide (H₂O₂) signaling in wheat plants. Wheat plants increase antioxidant defense mechanisms under abiotic stresses, such as drought, cold, heat, salinity and UV-B radiation, to alleviate oxidative damage. Moreover, H₂O₂ signaling is an important factor contributing to stress tolerance in cereals.     

Nutrient accumulation and translocation in maturing wheat plants grown on waterlogged soil

Wheat plants (Triticum aestivum L., cv. Arina) growing in large pots (perforated at the bottom for controls, intact for flooding) were embedded in the field in spring. Waterlogging was initiated at anthesis and was maintained throughout the maturation period. Grain yield as well as potassium, phosphorus and magnesium contents in the shoot were decreased on flooded soil, while manganese and iron contents increased considerably. Total calcium and zinc contents per shoot remained comparable to those in controls. The reduction of potassium, phosphorus and magnesium contents by waterlogging was greatest in the grains, while manganese and iron accumulated mostly in the vegetative parts and the glumes. Zinc contents were also lowered in the grains during waterlogging due to an inhibited redistribution from the vegetative parts to the grains. Our results indicate that flooding caused not only an accumulation of manganese and iron in the shoot, but also affected the redistribution of macro- and micronutrients to the maturing gains.     

Characterization of photosynthetic pigment composition, photosystem II photochemistry and thermal energy dissipation during leaf senescence of wheat plants grown in the field

Photosynthetic pigment composition and photosystem II (PSII) photochemistry were characterized during the flag leaf senescence of wheat plants grown in the field. During leaf senescence, neoxanthin and beta-carotene decreased concomitantly with chlorophyll, whereas lutein and xanthophyll cycle pigments were less affected, leading to increases in lutein/chlorophyll and xanthophyll cycle pigments/chlorophyll ratios. The chlorophyll a/b ratio also increased. With the progression of senescence, the maximal efficiency of PSII photochemistry decreased only slightly in the early morning (low light conditions), but substantially at midday (high light conditions). Actual PSII efficiency, photochemical quenching and the efficiency of excitation capture by open PSII centres decreased significantly both early in the morning and at midday and such decreases were much greater at midday than in the early morning. At the same time, non-photochemical quenching, zeaxanthin and antheraxanthin contents at the expense of violaxanthin increased both early in the morning and at midday, with a greater increase at midday. The results in the present study suggest that a down-regulation of PSII occurred in senescent leaves and that the xanthophyll cycle plays a role in the protection of PSII from photoinhibitory damage in senescent leaves by dissipating excess excitation energy, particularly when exposed to high light.     

Different photosynthetic responses of wild and cultivated plants to high irradiance

In addition to other factors, high altitude (HA) environment is characterized by high photosynthetic photon flux density (PPFD). Photosynthetic characteristics of wild and cultivated plants were studied at different irradiances at Losar, India (altitude 4 200 m). Wild plants were tolerant to high PPFDs. Slopes of curve between net photosynthetic rate (P _N) and intercellular CO₂ concentration (C _i) or stomatal conductance (g _s) increased with increase in irradiance suggesting insensitivity or tolerance of these plants to higher PPFD. Cultivated plants, however, were sensitive to higher PPFD, their slopes of curves between P _N and C _i or g _s decreased with increased PPFD. Tolerance or insensitivity to higher PPFD was an important parameter affecting plant performance at HA.     

Effects of growth temperature on photosynthetic carbon metabolism in green plants III. Differences in structure, photosynthetic activities and activities of ribulose diphosphate carboxylase and glycolate oxidase in leaves of wheat grown under varied temperatures

The rate of ferricyanide photoreduction in broken chloroplastsisolated from leaves of wheat acclimatized to a low temperature(mean temperature, 5–7?C) was similar to that in chloroplastsfrom wheat acclimatized to a high temperature (20–25?C). There was no practical difference in glycolate oxidase activityin leaf extracts of wheat plants grown at low and high temperatures.In contrast, the ribulose diphosphate carboxylase activity ofchloroplasts from low temperature sample was less than halfthat for the high temperature sample. Chloroplasts having a high rate of photosynthetic CO₂-fixationwere obtained from wheat acclimatized to a low temperature,whereas the CO₂-fixation activity in chloroplasts isolated fromhigh temperature-acclimatized wheat was very low. Electron microscopy revealed that chloroplasts in high temperature-acclimatizedwheat were ellipsoidal, electron dense and contained starchgranules. Those in low temperature-acclimatized leaves wereround and did not contain starch granule. ²Present address: Department of Botany, Faculty of Science,University of Tokyo, Tokyo, Japan (Received August 7, 1973; )     

Age-dependent changes of photosynthetic responses induced by electrical signals in wheat seedlings

Electrical signals in plants, namely, the action potential (AP) and variation potential (VP) alter the activity of many processes, including photosynthesis. The functional responses induced by electrical signals vary in direction and amplitude, which might be determined by variable conditions of plants prior to stimulation, by the development stage in particular. In this work, the parameters of VP-induced photosynthetic responses were analyzed at various stages of wheat seedling development. Local wounding of the second leaf in wheat plants induced the propagation of VP and altered the activity of photosynthesis at a distance from the wound location. The amplitude of VP was enlarged when the seedling age increased from 11 to 18 days. The VP-induced photosynthetic response changed with age both qualitatively and quantitatively. The amplitude of VP-induced changes in CO₂ assimilation and nonphotochemical quenching (NPQ) increased with age, which might be due to the increase in VP amplitude and associated changes in Ca²⁺ and H⁺ concentrations. The quantum yield of photosystem II photoreaction was subject to age-dependent changes: the photochemical quantum yield (γ(PSII)) was found to increase after VP in young leaves, whereas the decline in γ(PSII) was observed after the VP propagation in mature leaves. The results may explain the diversity of photosynthetic responses caused by the electrical signals.     

Senescence and protein remobilisation in leaves of maturing wheat plants grown on waterlogged soil

Nicotianamine (NA), the key precursor of the mugineic acid family phytosiderophores (MAs), is synthesized from S-adenosylmethionine (SAM). The NA synthase was strongly induced by Fe-deficiency treatment, and the activity increased to the maximum level faster than the time of maximum level of MAs secretion and also before the appearance of severest chlorosis. The enzyme was mainly localized in the roots of barley. NA synthase had the optimum pH at 9.0, a molecular weight of about 40,00050,000 estimated by gel filtration or about 30,000 by SDS-PAGE. Using hydrophobic chromatography, hydroxylapatite chromatography, and preparative SDS-PAGE, NA synthase was purified as one band on SDS-PAGE.     

Structural and functional reorganization of the photosynthetic apparatus in adaptation to cold of wheat plants

In seedlings of a cold-resistant wheat variety, the dynamics was studied of the main structural-functional parameters of the photosynthetic apparatus (PSA) and of cold resistance of leaf cells in low-temperature plant adaptation. It has been established that a complex of structural-functional PSA changes takes place in seedling leaves under the influence of cold. As a result, as early as in the first hours of hardening, the formation of chloroplasts begins to occur in mesophyll cells of larger sizes and with a thylakoid system of the “sun type.” Owing to structural and functional readjustment (a change of content of pigments, stabilization of pigment-protein complexes, and enhancement of nonphotochemical quenching of excess energy) in the process of cold adaptation, the rate of photosynthesis stabilizes. It is suggested that the observed structural-functional PSA rearrangement is a necessary condition for formation of increased cold resistance of leaf cells; this, alongside with other physiological-biochemical changes occurring in parallel in cells and tissues of the plants, provides their survival under conditions of low temperatures.     

From fresh vegetables to the harvest of wheat plants grown in the "SVET" space greenhouse onboard the MIR orbital station

Researchers report the results of experiments conducted onboard MIR in 1990, 1995, and 1996 in raising edible crops. In the 1990 experiment, radishes and Chinese cabbage were grown successfully, though the experimental plants were up to four times smaller than controls at harvest. The 1995 experiment in growing wheat through a complete life cycle was not completed. The 1996 experiment was successful in growing wheat through a complete life cycle to the seed stage. No seeds developed on any of the 279 ears harvested in that experiment. Reasons for the seedless development are explored.     

Reactive oxygen intermediates produced by photosynthetic electron transport are enhanced in short-day grown plants

Leaves of tobacco plants grown in short days (8h light) generate more reactive oxygen species in the light than leaves of plants grown in long days (16h light). A two fold higher level of superoxide production was observed even in isolated thylakoids from short day plants. By using specific inhibitors of photosystem II and of the cytochrome b(6)f complex, the site of O(2) reduction could be assigned to photosystem I. The higher rate of O(2) reduction led to the formation of a higher proton gradient in thylakoids from short day plants. In the presence of an uncoupler, the differences in O(2) reduction between thylakoids from short day and long day plants were abolished. The pigment content and the protein content of the major protein complexes of the photosynthetic electron transport chain were unaffected by the growth condition. Addition of NADPH, but not of NADH, to coupled thylakoids from long day plants raised the level of superoxide production to the same level as observed in thylakoids from short day plants. The hypothesis is put forward that the binding of an unknown protein permits the higher rate of pseudocyclic electron flow in thylakoids from short-day grown plants and that this putative protein plays an important role in changing the proportions of linear, cyclic and pseudocyclic electron transport in favour of pseudocyclic electron transport. This article is part of a Special Issue entitled: Photosynthesis Research for Sustainability: from Natural to Articifical.     

Dependence of photosynthetic rates on leaf density thickness in deciduous woody plants grown in sun and shade

Comparisons of photosynthetic rates were made on leaves of ten species of woody dicotyledons grown in the field under full sun or under a canopy which transmitted approximately 18% of full light. Photosynthesis and dark respiration were measured and compared on various bases: area, chlorophyll, fresh weight of lamina, density thickness (fresh weight per unit area), and protein.

Light-saturated photosynthesis per unit area or unit chlorophyll was about 1.5 times greater in the sun leaves than in the shade leaves and essentially equal per unit fresh weight or unit protein. Sun leaves were thicker but the enzymes per unit fresh weight remained constant as thickness varied. Chlorophyll per unit area remained about constant; chlorophyll per unit fresh weight varied inversely with changes in leaf thickness. Thus, density thickness variation is important in photosynthetic adaptation to sun and shade. This is also shown by the relationship between light-saturated photosynthesis per unit area and density thickness.

     

Growth and photosynthetic responses in Jatropha curcas L. seedlings of different provenances to watering regimes

Seedlings from four provenances of Jatropha curcas were subjected to 80, 50, and 30% of soil field capacity in potted experiments in order to study their responses to water availability. Our results showed that with the decline of soil water availability, plant growth, biomass accumulation, net photosynthetic rate, stomatal conductance (g_s), and transpiration rate (E) decreased, whereas leaf carbon isotope composition (δ¹³C), leaf pigment contents, and stomatal limitation value increased, while maximal quantum yield of PSII photochemistry was not affected. Our findings proved that stomatal limitation to photosynthesis dominated in J. curcas under low water availability. The increase of δ¹³C should be attributed to the decrease in g_s and E under the lowest water supply. J. curcas could adapt to low water availability by adjusting its plant size, stomata closure, reduction of E, increasing δ¹³C, and leaf pigment contents. Moreover, effects of provenance and the interaction with the watering regime were detected in growth and many physiological parameters. The provenance from xeric habitats showed stronger plasticity in the plant size than that from other provenances under drought. The variations may be used as criteria for variety/provenance selection and improvement of J. curcas performance.     

Protective and injuring action of visible light on photosynthetic apparatus in wheat plants during hyperthermia treatment

Illumination of wheat (Triticum aestivum L.) leaves during heat treatment produced either additional injury or protection of photosynthetic apparatus depending on irradiance and the heating dose. Furthermore, illumination of leaves during hyperthermia exerted differential impacts on thermal tolerances of photosynthesis and photosystem II-driven electron transport assessed from the reduction of 2,6-dichlorophenolindophenol (DCPIP). Measurements with infrared gas analyzer showed that mild heating of leaves in darkness (10 min at 38–40°C) had stronger inhibitory effect on CO₂ uptake than heating of leaves exposed to low and moderate complex irradiances (3–30 klx), as well as excessive irradiance (75–100 klx). When the leaves were heated at higher temperatures (42–44°C), the low and moderate irradiances had a protective action, while high-intensity light aggravated the inhibition of photosynthesis. Illumination of leaves with weak light during heat treatment mitigated the impairment of chloroplast ultrastructure, whereas irradiation with high-intensity light (100 klx) destroyed the sensitive population of chloroplasts. The heat-stimulated photoinhibition was stronger for leaf photosynthesis than for DCPIP reduction in chloroplasts isolated from heat-treated leaves. No correlation was observed between the extent of violaxanthin deepoxidation, zeaxanthin accumulation, and the protective effect of light on photosynthetic apparatus during heat treatments.     

Growth and photosynthetic responses to salinity of the salt-marsh shrub Atriplex portulacoides

BACKGROUND AND AIMS: Atriplex (Halimione) portulacoides is a halophytic, C(3) shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology. METHODS: The responses of young plants to salinity in the range 0-700 mol m(-3) NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined. KEY RESULTS: Plants accumulated Na(+) approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m(-3) NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO(2) assimilation, which were not affected by 200 mol m(-3) but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO(2) concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Phi(PSII)) nor the chlorophyll fluorescence ratio (F(v)/F(m)) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F(v)/F(m) was in fact depressed more at low external sodium concentration, by the end of the experiment. CONCLUSIONS: The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.     

Growth and photosynthetic responses of four landscape shrub species to elevated ozone

Attention should be paid to ozone (O₃) sensitivity of greening plant since ground-level O₃ concentrations are increasing especially in urban and suburban area. We studied the ecophysiological responses to elevated O₃ of four shrub species [Euonymus bungeanus Maxim. (EB), Photinia × fraseri (PF), Chionanthus retusus Lindl. & Paxt. (CR) and Cornus alba L. (CA)], which are often used for garden greening in China. Saplings of those species were exposed to high O₃ concentration (70 nmol mol⁻¹, 7 h d⁻¹ for 65 d) in open-top growth chambers. Responses to O₃ were assessed by gas exchanges, chlorophyll (Chl) fluorescence and dry mass. We found that elevated O₃ significantly decreased lightsaturated net photosynthetic rate (P _Nsat), transpiration rate (E) and stomatal conductance (g _s). The ratio of intercellular CO₂ to ambient CO₂ concentration (C _i/C _a) did not reduce under O₃ fumigation which suggested that the O₃-induced depressions of P _Nsat under O₃ fumigation were probably due to limitation of mesophyll processes rather than stomatal limitation. High O₃ exposure also significantly depressed the maximum efficiency of photosystem II (PSII) photochemistry in the dark-adapted state (F_v/F_m) which meant the O3-induced photoinhibition. Both root dry mass and root/shoot ratios were significantly decreased under ozone fumigation, but the total mass was unchanged. The responses of gas exchange such as P _Nsat in these four shrubs to O₃ exposure were species-specific. Highest loss of P _Nsat was observed in EB (−49.6%), while the CR had the lowest loss (−36.5%). Moreover, the O₃-exposed CR showed similar g _s as CF, reflecting that its O₃ flux might be unchanged under elevated O₃ environment. Ozone drastically decreased actual quantum yield of PSII (Φ_PSII) and electron transport rate (ETR) in EB while increased Φ_PSII and ETR in CR. Furthermore, the relative losses in P _Nsat positively correlated with the relative decreases in Φ_PSII and ETR which indicated that the impairment of photosynthesis was probably affected by the light reaction process. The light reaction of EB was impaired most seriously but that of CR was not damaged. All results indicated that EB was probably the most sensitive shrub species to O₃ while CR the most tolerant one. Therefore, CR might be an ideal choice for greening in ozone-polluted areas.